Revision of the Central Asian Anomalobuthus Kraepelin, 1900, with Descriptions of Three New and a Generic Synonymy (Scorpiones: )

Rolando Teruel, František Kovařík & Victor Fet

October 2018 – No. 270 Occasional Publications in Scorpiology

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Publication date: 23 October 2018

http://zoobank.org/urn:lsid:zoobank.org:pub:9D0AA0A5-38D9-49DB-B2FB-380550213399 Euscorpius — Occasional Publications in Scorpiology. 2018, No. 270

Revision of the Central Asian scorpion genus Anomalobuthus Kraepelin, 1900, with descriptions of three new species and a generic synonymy (Scorpiones: Buthidae)

Rolando Teruel 1, 2, František Kovařík 3 & Victor Fet 4

1 Grupo de Sistemática y Ecología de Artrópodos Caribeños, Calle 200 # 3759, e/ 37 y 45, Reparto Versalles, La Lisa, La Habana 13500, Cuba; [email protected] 2 Instituto de Ecología y Sistemática (Subdirección de Colecciones Zoológicas), Carretera de Varona # 11835, e/ Oriente y Lindero, Reparto Calabazar, Boyeros, La Habana 11900, Cuba 3 P. O. Box 27, CZ - 145 01 Praha 45, Czech Republic; www.scorpio.cz 4 Department of Biological Sciences, Marshall University, Huntington, WV, 25755, USA; [email protected]

http://zoobank.org/urn:lsid:zoobank.org:pub:9D0AA0A5-38D9-49DB-B2FB-380550213399

Summary

We revise the Central Asian endemic genus Anomalobuthus Kraepelin, 1900, which was considered monotypic for more than 100 years until the recent addition of a second species from Iran (Teruel et al., 2014). We redefine the generic diagnosis of Anomalobuthus and reveal that it is composed of no less than six species, three of which are described as new: A. krivochatskyi, sp. n. (central Uzbekistan and extreme southern Kazakhstan), A. lowei, sp. n. (southeastern Kazakhstan), and A. pavlovskyi, sp. n. (south-central Kazakhstan and extreme northern Turkmenistan). The monotypic genus Psammobuthus Birula, 1911 (described from the Ferghana Valley at the border of Uzbekistan and Tajikistan) is synonymized under Anomalobuthus; its single species is transferred as Anomalobuthus zarudnyi (Birula, 1911), comb. n. We also clarify the type locality of the type species A. rickmersi Kraepelin, 1900, from accurate data published by its collector Rickmer Rickmers (1913) but overlooked since its very original description: it is Baljuvon in Tajikistan. Occurrence of A. rickmersi in southern and southeastern Turkmenistan is confirmed. All available species are fully illustrated with color photos of habitus and morphologically diagnostic characters, and a key covering all six recognized species of Anomalobuthus is included.

Introduction del based on DNA markers, including A. rickmersi and a co-distributed convergent psammophile Liobuthus A rare genus Anomalobuthus (Scorpiones: Buthi- kessleri Birula, 1898, was offered by Graham et al. dae) was described by Karl Kraepelin (1900) with its (2012) who also noted a phylogenetic structure between type species A. rickmersi based on a single type spec- known populations of Anomalobuthus. imen, currently housed at Zoologisches Museum Ham- The genus Anomalobuthus remained monotypic burg, Germany (below, ZMUH). The specimen was until the recent discovery of a second, new species in captured in “Bucharei”, i.e., the Emirate of Bukhara eastern Iran, A. talebii (Teruel, et al., 2014). Further (then a protectorate of the Russian Empire) by the revisionary work of our research group has revealed that famous German explorer Willi Rickmer Rickmers the populations assigned in literature to “A. rickmersi” in (1873–1965) who visited Central Asia several times. fact include at least four species, three of which are Later, Anomalobuthus rickmersi Kraepelin, 1900 was described below as new ones. Our study shows the pres- recorded in many deserts of Central Asia (Fet, 1989). ence of at least six species in the genus Anomalobuthus, Sand desert scorpions of Central Asia, including A. found in five countries: Iran (one species), Kazakhstan rickmersi, were discussed by Fet et al. (1998) who noted (three species), Tajikistan (first record; two species), high parallelism in the adaptive features allowing psam- Turkmenistan (two species), and Uzbekistan (two mophily (life in sand). Recently, a phylogeographic mo- species). 2 Euscorpius — 2018, No. 270

Methods & Material Psammobuthus Birula, 1911a: 69–74; Werner, 1934: 271; Fet, 1989: 119; Sissom, 1990: 50, 52; Fet & Nomenclature and measurements follow Stahnke Lowe, 2000: 214 (includes full list of references (1971), Kovařík (2009), and Kovařík & Ojanguren Af- before 1998); Fet et al., 2001: 183, 185, tab. 1. New filastro (2013), except for trichobothriotaxy (Vachon, synonym. 1974), metasomal carinae (Francke, 1977), pedipalp chela carinae (Acosta et al., 2008, as interpreted by Ar- DIAGNOSIS. Adult size small for the family (males mas et al., 2011), and sternum (Soleglad & Fet, 2003a). 22–34 mm, females 23–37 mm). Adults slightly dimor- In the text, the redescription of the type species is phic: males smaller and slenderer, with pectines larger presented first and the descriptions of the three new and with higher tooth counts. Cheliceral dentition species follow ordered alphabetically; a shorter des- modified from most common buthid pattern (Vachon, cription of the species transferred to this genus is given 1963): both fixed and movable finger ventrally with a last, taken from the original description because the single denticle. Pedipalps very slender, with chelae nar- types (only known specimens) were not available for rower than patella; trichobothrial pattern A-β, varying study. from apparently orthobothriotaxic to clearly neobothrio- Scanning Electron Microscopy was performed by taxic (femoral d2 and chelal Et, V1, and esb trichobothria Jan Štundl at Charles University, Prague, Czech Repub- absent to rudimentary), patellar trichobothrium d3 lo- lic, and Victor Fet (with the expert help of David Neff, cated between dorsomedian and dorsointernal carinae; Kelly Anne Daniel, and Jacqueline Aisling) at Marshall fingers without lobe/notch combination, with 8–11 University, West Virginia, USA. principal rows of denticles (division of basalmost rows Specimens studied herein are preserved in 80% usually poorly defined), all arranged in a straight line, ethanol and deposited in the following collections: external accessory denticles entirely absent, internal Naturhistorisches Museum Wien, Vienna, Austria accessory denticles absent from basalmost rows, mov- (NHMW); National Museum of Natural History, Prague, able finger with 1–2 accessory denticles basal to the Czech Republic (NMPC); National Museum of Natural terminal denticle. Carapace essentially without carinae, History, Bulgarian Academy of Sciences, Sofia, Bulgar- with anterior margin convex; median eyes very large and ia (SOFM); Zoological Institute, Russian Academy of raised, five pairs of much smaller lateral eyes (three Sciences, St. Petersburg, Russia (ZISP); Zoological same-sized and aligned along each anterolateral corner, Museum, Moscow State University, Moscow, Russia plus two tiny and offset above the former). Tergites I–VI (ZMMSU); Centrum für Naturkunde (CeNak); Center of vestigially to obsoletely tricarinate. Sternum type 1, Natural History Universität Hamburg, Zoological Mus- relatively small, and triangular to subpentagonal in eum, Germany (ZMUH); Zoologische Sammlung der shape; posterior depression very large and deep. Pectines Universität, Rostock, Germany (ZSRO); and personal very large, with 22–27 teeth in males and 19–26 in collections of Rolando Teruel (RTOC), František females; fulcra well developed; lamellae and basal plate Kovařík (FKCP), Victor Fet (VFPC), and Graeme Lowe unmodified. Legs very long and slender, adapted to (GLPC). psammophily: tibia and tarsi of I–III short, curved, flat, and paddle-like, with setation heavily modified into Systematics bristlecombs, telotarsi with two rows of long setae on ventral surface, claws very long, asymmetrical, and Family Buthidae C. L. Koch, 1837 weakly curved; tibial spurs highly variable, from fully developed to entirely absent (usually highly reduced), Genus Anomalobuthus Kraepelin, 1900 tarsal spurs well-developed but with complex armature Figures 1–141 (bifurcate, with setae and spines). Sternites with spir- acles slit-like; V without a well-defined smooth patch. Anomalobuthus Kraepelin, 1900: 1–11, figs. 4–7; Birula, Metasoma slender, with carination highly reduced; 1917: 55; Werner, 1934: 272; Vachon, 1974: 908, dorsal lateral and lateral supramedian carinae of fig. 37; Fet, 1989: 80; Sissom, 1990: 37, 46, 48–51, segments I–IV with 1–2 conspicuously enlarged pos- 101, 120, figs. 3.17h, m; Fet & Lowe, 2000: 75 terior terminal denticles; ventral lateral carinae of seg- (includes full list of references before 1998); Capes ment V evenly flared, with sharp to round denticles; & Fet, 2001: 300–301; Fet et al., 2001: 183–185, intercarinal spaces with some coarse punctuations on tab. 1; Fet et al., 2003: 4; Fet et al., 2005: 2–3, 6–7, IV–V, each surrounding the base of a macroseta and 10–11, 13, 22, 24–25, figs. 10, 23–25, tab. 1; usually along carinae. Telson elongate, subaculear Lourenço, 2005: 111; Volschenk et al., 2008: 652, tubercle absent. 654, 659–660, Fig. 1I; Graham et al., 2012: 96; Teruel et al., 2014: 1–10, figs. 1–24, tabs. 1–2. Nec DISTRIBUTION (Fig. 141): Iran, Kazakhstan, Tajik- Lourenço, 2001: 15–20 [misidentification]. istan, Turkmenistan, and Uzbekistan.

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 3

Description of several distinct species of Anomalo- holotype and apparently not even a member of Anomalo- buthus attests to differentiation of this psammophile buthus (see below, in the Notes section of that species). genus in isolated pockets of Central Asian sand deserts, a biogeographic process well documented in other SUBORDINATE TAXA. Anomalobuthus rickmersi Kraepe- groups (see below in Biogeography section). The lin, 1900 (Turkmenistan, Tajikistan), A. krivochatskyi genus Anomalobuthus is found from eastern Iran, at Teruel, Kovařík et Fet, sp. n. (Uzbekistan, Kazakhstan), 33°20'49.45"N, to Kazakhstan, at 44°20'37"N (Baiga- A. lowei Teruel, Kovařík et Fet, sp. n. (Kazakhstan), A. kum). Thus its range spans over 10 degrees (>1,100 km) pavlovskyi Teruel, Kovařík et Fet, sp. n. (Kazakhstan, from north to south across the great deserts of Central Turkmenistan), A. talebii Teruel, Kovařík, Navidpour et Asia and Iran. From west to east, it spans over 23 Fet, 2014 (Iran), A. zarudnyi (Birula, 1911), comb. n. degrees (>2,600 km) from Kheles, Turkmenistan, at (Uzbekistan, Tajikistan). 53°24'E, to Kapchagay, Kazakhstan, at 77°05'E (Teruel et al., 2014). Anomalobuthus rickmersi Kraepelin, 1900 The presence of Anomalobuthus in Afghanistan, Figures 1–33, 117–119, 126–127, 137, 141; Tables 1, 4–5 Kyrgyzstan, and China has never been documented, but it can be predicted to occur there in continuous or Anomalobuthus rickmersi Kraepelin, 1900: 1–11, similar habitats since at least four species occur very figs. 4–7; Birula, 1904: 32; Birula, 1905: 449 (in part); near to their borders: A. lowei sp. n., A.rickmersi, A. Birula, 1911a: 73–74 Birula, 1911b: 171 (in part); Rick- talebii, and A. zarudnyi comb. n. (Fig. 141). mer Rickmers, 1913: 442; Birula, 1917: 112 (in part);

Pavlovsky, 1934: 201 (in part). Werner, 1934: 272; fig. ECOLOGY. Almost all species are ultra-psammo- 341; Weidner, 1959: 98; Stahnke, 1972: 122 (in part), philes, which inhabit exclusively the vast sand dune fig. 3; Vachon, 1974: 908, fig. 37; Fet, 1980: 224 (in systems of the various Central Asian deserts. The single part); Fet, 1989: 80 (in part); Sissom, 1990: 37, 46, 48– exception is the type species A. rickmersi, a widespread 51, 96, 101, 120, figs. 3.17h, m; Fet, 1994: 531 (in part); psammophile that also reaches the upper clay taluses of Gromov & Kopdykbaev, 1994: 20 (in part); Fet et al., the Kyzylsu Valley. 1998: 613 (in part), fig. 6 [specimen from Repetek]; Fet

& Lowe, 2000: 75 (in part, includes full list of ref- NOTES. erences before 1998); Fet et al., 2001: 183–185, tab. 1 1. The genus Psammobuthus Birula, 1911, was des- (in part). Fet et al., 2003: 4 (in part); Fet et al., 2005: 2– cribed based on two specimens from the sands of the 3, 6–7, 10–11, 13, 22, 24–25, figs. 10, 23–25, tab. 1 (in Fergana Valley, in the boundary of modern Uzbekistan part); Graham et al., 2012: 95–96 (in part); Warburg, and Tajikistan. It was never collected again and Fet et al. 2013: 96, 102 (in part); Teruel et al., 2014: 1, 4, 6–7, 9, (2005: 13) quoted an earlier opinion of A. V. Gromov fig. 20, tabs. 1–2 (in part). Nec Lourenço, 2001: 15–20 (pers. comm., 2002) that it could be a junior synonym of [misidentification, see below in Remarks section]. Anomalobuthus. The present study of abundant material of the latter, confirmed this suspicion: all characters used to diagnose Psammobuthus by Birula (1911a) are dis- HOLOTYPE ♂ (ZMUH, examined). Tajikistan, tributed within the variability range now known for the Khatlon Province, Kyzylsu River valley, near Baljuvon entire genus. Thus, the following synonymy is estab- (= Bal'dzhuvon or Baljuan), early summer 1896, leg. W. lished herein: Anomalobuthus Kraepelin, 1900 = Rickmer Rickmers. See discussion below, in Remarks Psammobuthus Birula, 1911, new synonym. section.

2. Uzbekistan has been traditionally listed as a country OTHER MATERIAL EXAMINED. Turkmenistan, Kara- record for the genus Anomalobuthus, based on the in- kum Desert, Lebap (=Chardzhou) Province, Repetek correct assumption that the holotype of A. rickmersi Nature Reserve, 38°33'54"N 63°10'51"E, 201 m a.s.l., originated from this country (see below, in the section 21 July 1985, leg. V. Fet, 2♂♂ (NHMW), 5♂♂, 1 pertinent to this species). In fact, the first published juvenile (VFPC); same data except 29 April 1967, leg. specimens originating actually from Uzbekistan (modern V. I. Kuznetsov, 1♂ (ZSRO Sc-1249); same data except Bukhara Province) were collected only in 2002 by V. 15 April 2002, leg. V. Fet, 1 juvenile (VFPC); Karakum Fet and A. V. Gromov, and first studied by Soleglad & Desert, Akhal Province, Tejen (= Tedzhen, Tedschen), Fet (2003a, 2003b) and Fet et al. (2005). 37°22'39"N 60°29'52"E, 188 m a.s.l., 10–24 July 1904 [23 July–6 August], leg. K. Aris, 1♂ (ZISP 809); Mary 3. Lourenço (2001) published a “revised diagnosis” of Province, Serkhetabad District, Badghyz Plateau, Chai- the genus, allegedly based upon the holotype of A. nury, 35°40'52"N 62°01'34"E, 458 m a.s.l., 6 April rickmersi. However, it is entirely wrong because the 2002, leg. V. Fet & A. V. Gromov, 1♀ (FKCP), 1♀ specimen examined by Lourenço was not the actual (NMPC). 4 Euscorpius — 2018, No. 270

Figures 1–3: Anomalobuthus rickmersi, adult male holotype: full-body views, dorsal (1) and ventral (2), plus original labels (3). Scale bar = 10 mm (1–2). The red inset at bottom right shows the label of the specimen examined by Lourenço (2001), demonstrating it was not the actual Rickmers's type specimen.

DIAGNOSIS (emended). Adult size 30–34 mm in accessory denticles. Pectines with 23–27 teeth in males males, 32–37 mm in females. Coloration light yellow, and 20–23 in females. Tibial spur variable from absent essentially immaculate, only with some regular blackish (specimens from Turkmenistan) to present but very spots on pedipalps and metasoma (darker and more small on leg IV only (male holotype from Tajikistan). extended in female); metasomal segment V and telson Metasoma conspicuously attenuate, with most carinae reddish brown (male) to black (female). Pedipalp fingers obsolete to vestigial; ventral lateral carinae of segment V with 9–10 principal rows of denticles and 7–9 internal composed of denticles narrowly conical (male) to nar-

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 5

Figures 4–7: Anomalobuthus rickmersi from Turkmenistan: adult male from Repetek (4–5) and adult female from Chainury (6– 7), full-body views, dorsal (4, 6) and ventral (5, 7). Scale bar = 10 mm. rowly lobate (female), dorsal lateral and lateral supra- icle enlarged; intercarinal spaces essentially smooth on median carinae of segments I–IV with terminal dent- segments I–III, sparsely granulose on ventral and lat-

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Figures 8–13: Anomalobuthus rickmersi. Figures 8–9. Juvenile male from Repetek, Turkmenistan: full-body views, dorsal (8) and ventral (9). Scale bar = 10 mm. Figures 10–13. Adult male holotype: chelicerae, carapace and tergites I–III, dorsal (10), sternopectinal region and sternites III–V, ventral (11), distal part of left leg III, internal (12), distal part of left leg IV, internal (13). eral areas of IV and very densely granulose on ventral lowish; in general, the base color is paler on pedipalp and lateral areas of V (granulation sparser in female). chelae, legs, and pectines. Chelicerae immaculate, ex- Telson vesicle elongate oval and sparsely setose. cept for dark brown finger teeth. Pedipalp femur and patella with internal surface infuscate; chela immaculate, REDESCRIPTION (adult male holotype). Coloration only with finger denticles dark brown. Carapace im- (Figs. 1–2) moderately faded and translucent due to long maculate, only with a blackish spot under every ocular preservation, but still reliably traceable. Base light yel- group. Tergites immaculate. Coxosternal region and

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 7 genital operculum immaculate. Pectines pale yellowish, Pectines (Fig. 11). Standard-sized for the genus: immaculate. Sternites immaculate. Legs essentially im- very long, extending beyond leg IV coxa-trochanter maculate; claws with distal half dark brown. Metasoma joint), subrectangular and densely setose. Tooth count not conspicuously bicolor, only becoming progressively 25/25. Basal plate heavily sclerotized, much wider than darker and redder distally, segments I–IV with a dif- long, anterior margin with a very deep, narrow antero- fusely annulated appearance (basal and distal parts of median furrow, posterior margin very shallowly convex. each segment faintly infuscate as thin brown rings, Legs (Figs. 12–13). Very slender, with all carinae which become larger and more diffuse dorsally) and weak and subgranulose to subcostate; intercarinal tegu- segment V faintly infuscate, becoming progressively ment smooth and glossy. Tibial spurs entirely absent on darker and redder distally; carinae not infuscate nor un- legs III, reduced to a sinuous remnant on legs IV. derlined with dark pigment. Telson vesicle reddish Mesosoma (Figs. 10–11). Tergites with the same brown, aculeus with basal half yellowish and distal half sculpture as on carapace; I–VI irregularly tricarinate: the dark brown. median longitudinal carina is weak, short, and formed by Chelicerae (Fig. 10). With dentition typical for the irregular medium-sized granules that do not project be- genus, as described for A. krivochatskyi sp. n. (see yond posterior margin, but the submedian carinae are below). defined only by few coarse granules irregularly aligned; Pedipalps (Figs. 14–22). Relatively short but very tergite VII with five well-defined carinae: the median slender, essentially bare. Femur subtly curved inwards, carina is very short but strongly subserrate, the sub- with carinae weak, granulose to subdenticulate; inter- medians and laterals are long, strong and finely serrate carinal tegument smooth and glossy. Patella straight, to denticulate. Sternites sparsely setose; III–VI glossy with carinae obsolete to absent, smooth except on and with very subtle vestiges of smooth submedian internal surface where vestigially denticulate; inter- carinae, spiracles relatively short and slit-like, almost carinal tegument smooth and glossy. Chela elongate and transversely arranged (not strongly oblique), V with very slender; manus conspicuously narrower than patella smooth patch absent; VII with two pairs of carinae: the (ratio 0.79), subcylindrical (1.80 times longer than wide, submedians are long and finely crenulate to subcostate 1.07 times wider than deep), with carinae weak to mod- and the laterals are very short and subcrenulate, inter- erate, smooth; intercarinal tegument smooth and glossy; carinal tegument coriaceous to minutely granulose. fingers very long (movable finger 2.44 times longer than Metasoma (Figs. 23–25). Elongated and slightly underhand), only subtly curved and with 10 principal wider subdistally; with 10/10/10/8/5 complete to almost rows of denticles (the two basalmost rows are complete carinae, almost all obsolete to vestigial and moderately well-defined), basal lobe/notch combination smooth, with coarse punctations scattered: dorsal laterals absent, external accessory denticles absent, internal very weak on I–III (with one terminal denticle enlarged), accessory denticles very large and claw-like (increasing obsolete to vestigial as rounded ridges on IV, absent on in size distally), numbering eight on both fixed and V; lateral supramedians very weak on I–III (with one movable fingers, movable finger with one claw-like terminal denticle enlarged), obsolete to vestigial as accessory denticle basal to the very large terminal rounded ridges on IV–V; lateral inframedians obsolete denticle. on I–III, absent on IV–V; ventral laterals obsolete on I– Carapace (Fig. 10). Very strongly trapezoidal IV, weak to moderate on V, where become progressively (much narrower anteriorly) and wider than long; anterior stronger and somewhat flared distally, formed by margin shallowly convex, with 5–6 pairs of thin macro- sharply serrate, subequal denticles; ventral submedians setae and some very short microsetae. Carination essen- obsolete on I–IV, absent on V; ventral median absent on tially absent: the only carinae present are the super- I–IV, weak to moderate on V. Intercarinal tegument ciliaries, which are smooth. Furrows: anterior median, smooth and glossy, with minute but sharp granulation on median ocular, central median, posterior median and lateral and ventral surfaces, which become much denser posterior marginal fused, wide and moderately deep; towards distal segments. Dorsal furrow obsolete on all lateral oculars, lateral centrals, central transverse, and segments. Setation sparse, mostly represented by 5–8 posterior laterals long, narrow and shallow. Tegument dark macrosetae over every carina. very finely and densely granulose, with a few small to Telson (Figs. 26). Sparsely setose, with a few setae medium-sized granules scattered all over. scattered on dorsal and lateral surfaces. Vesicle elongate Sternum (Fig. 11). Standard for the genus, rel- oval (2.50 times longer than wide, 0.95 times wider than atively small and widely triangular in shape. deep), tegument smooth and glossy, with vestiges of Genital operculum (Fig. 11). Slightly damaged. coarse granules arranged into three obsolete longitudinal Relatively large, each half roundly subtriangular in carinae (ventral median and ventral submedians), and shape. Genital papillae present. ventrally with some coarse punctations. Subaculear tub-

8 Euscorpius — 2018, No. 270

Figures 14–26: Anomalobuthus rickmersi, adult male holotype: right pedipalp trochanter and femur, dorsal (14) and ventral (15), right pedipalp patella, dorsal (16), external (17) and ventral (18), right pedipalp chela, dorsal (19–20), external (21) and ventrointernal (22), metasoma and telson, lateral (23), dorsal (24) and ventral (25), telson, lateral (26). ercle absent. Aculeus conspicuously shorter than ves- darker and much more sharply patterned, e.g., pedipalp icle, thick and shallowly curved. femur with a conspicuous blackish spot on dorsodistal apex, which diffusely continues as a broad stripe over FEMALE (Chainury, Turkmenistan). Very similar to almost all internal surface, patella with internal surface male, sexual dimorphism evident by: 1) size com- blackish, carapace with anterior margin deeply infuscate, paratively larger inside each size-class; 2) coloration metasomal segment V and telson black; 3) mesosoma

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 9

Figures 27–33: Anomalobuthus rickmersi from Turkmenistan, adult male from Repetek (27–30) and adult female from Chainury (31–33): metasoma and telson, lateral (27, 31), dorsal (28, 32) and ventral (29, 33), telson, lateral (30). and metasoma slightly less slender; 4) genital papillae character alone is inappropriate to separate both pop- absent; 5) pectines smaller, with consistently lower tooth ulations taxonomically, especially when only a single counts; 6) metasomal segment V with lateral intercarinal specimen from Tajikistan is available. In addition, the tegument less granulose. See Figs. 6–7, 31–33 and Tabs. male from Tejen (ZISP 809) has tibial spurs on both legs 1, 4–5. IV as noted already by Birula (1905: 450)

VARIATION. Adult size varied from 30.4–34.1 mm COMPARISON. Adults of A. rickmersi can be very in males and 32.1–36.5 mm in females. easily distinguished by the greater slenderness of the Pectinal tooth count varied as follows: 23–27 teeth metasoma, especially in males (Figs. 1–2, 4–7, 117; in males and 20–23 in females, without clearly defined Tabs. 1, 4); even to unaided eye, most other congeners mode in either sex (Tab. 5). are more robust (Figs. 34–45, 65–66, 79–82, 90–91, 107, Males from Repetek are virtually identical to the 110, 113–114; Tabs. 2–4). For example, see the fol- holotype in all diagnostic characters. The only exception lowing male ratios: metasoma + telson length / carapace is that the tibial spurs are usually entirely absent. This length = 7.76 vs. 6.26–7.14, metasomal segments length 10 Euscorpius — 2018, No. 270

/ width = 1.74 vs. 1.02–1.35 (I), 2.13 vs. 1.35–1.82 (II), By the way, the “poisonous spider” mentioned by 2.20 vs. 1.39–1.88 (III), 2.56 vs. 1.75–2.23 (IV) and 2.71 the impressed collector is a solpugid, also described by vs. 1.85–2.38 (V). Kraepelin (1899). It is a member of the Gylippidae fam- Moreover, this species is the only one that has meta- ily, currently regarded as a junior synonym of Gylippus somal segment V and telson light reddish-colored in ferganensis Birula, 1893 (see Harms & Dupérré, 2018). adult males (Figs. 1–2, 4–5, 23–25, 27–29, 117); in all As all solpugids, it is indeed “vicious” but not “poison- other Anomalobuthus spp., it is blackish, entirely or at ous”. least in its distal half (Figs. 34–35, 38–39, 59–61, 79–80, Baljuan (also spelled Baldzhuan, now Baljuvon or 84–86). Bal'dzhuvon) is a well-known town in modern Tajik- istan, in the valley of Kyzylsu River, a tributary of the DISTRIBUTION (Fig. 141). Widespread in southern larger Panj River (= Pyandzh, Piandj), itself a tributary Central Asia, from the sands of the Karakum Desert of the great Amudarya (the ancient Oxus). Baljuvon (eastern Turkmenistan), to the loess soils of the Kyzylsu stands on the Silk Road, not far from the ancient Bactra River valley (southwestern Tajikistan). See further de- (now Balkh in northern Afghanistan), and was last time tails below, in Remarks section. noted as the place where Enver Pasha, a rival of the Turkish leader Mustafa Kemal (Ataturk), was killed in 1922 in a skirmish with the Soviet troops. ECOLOGY. According to the personal data of the In more peaceful times, Rickmer-Rickmers (1899) collector (Rickmer Rickmers, 1913: 442), the holotype published a map of his fourth travel to the Bukhara was collected in early summer, from crevices or cracks Emirate, conducted in 1898 (the 1896 trip was his third of fine alluvial soil (“loess crannies”). Populations from travel). We can see that he followed the same and (then) the Karakum Desert inhabit sand dunes but tend not to only route from Dushanbe (now the capital of Tajik- occupy loose sands where two other sympatric ultra- istan) to Khovaling, via Baljuvon. Baljuvon lies very far psammophiles are found, Liobuthus kessleri and to the southeast from the former Emirate's capital city of Pectinibuthus birulai Fet, 1984. Detailed ecological in- Bukhara (modern Uzbekistan) and is isolated by several formation from Repetek was published by Fet (1980). mountain ranges from the lowland deserts of Uzbek- istan. NOTES. The holotype of A. rickmersi remains the Although quite active zoological research was single specimen known from Tajikistan. Its collecting conducted in Central Asia by numerous Russian (and site was reported by Kraepelin (1900) as Bukhara then Soviet) researchers, very few scorpions have been (“Bucharei”, see the original museum label in Fig. 3) collected in the remote valleys of northern tributaries of and interpreted by all subsequent authors as originating Amu Darya in modern Uzbekistan and Tajikistan. It is from the city, the capital of the homonymous Bukhara not surprising, therefore, that no other specimens of A. Khanate, located in modern Uzbekistan. In addition, rickmersi have been collected in this area since 1896; in Birula (1911b: 171) erroneously stated that the type fact, no other scorpion collections we know of originate specimen of A. rickmersi originated from “westlichen from Baljuvon. Moreover, another, much more amazing Buchara” (i.e. western part of the Bukhara Emirate). scorpion from the same area was not discovered until the This specimen was collected by the famous German 1990s: the relict Pseudochactas ovchinnikovi Gromov, mountain explorer Willi Rickmer Rickmers (1873–1965) 1998 (Pseudochactidae). Its type locality lies in the who repeatedly visited the Bukhara Emirate, then a Surkhandarya River valley, which separates modern protectorate of the Russian Empire, which included part Uzbekistan from Tajikistan (Gromov, 1998; Fet et al., of modern Uzbekistan and Tajikistan (Rickmer-Rick- 2004). mers, 1899; Rickmer-Rickmers, 1913). During prep- The first known specimen from Turkmenistan (then aration of the present revision, we discovered that the Transcaspian Region of the Russian Empire) was Rickmer Rickmers (1913: 442) himself actually pub- reported by Birula (1904: 32, 35). It was a female lished unequivocal information on the precise collection collected next to the railway station of Repetek in the locality and date of the holotype, which we reproduce East Karakum, the future site of the famed Repetek below verbatim: Natural Reserve, by the coleopterologist Eduard Niko- laevich Fischer in February 1904 (ZISP 808). The “On the way to Baljuan I noticed that many poplar second specimen was also published by Birula (1905: trees were pitted with wormholes bored by that 449–450); it was a male from Tejen (=Tedzhen, stately Capricorn beetle the Pachydissus Sartus. In the early summer of 1896 the loess crannies around Tedshen) in Akhal Province, collected by K. Aris in had yielded up to my collection new specimens of June 1904 (ZISP 809). Further collections from Turk- poisonous spider and scorpion, Gylippus Rickmersi menistan published by Fet (1989), included a series of and Anomalobuthus Rickmersi. I feel highly gratified more than 20 specimens from Repetek (ZISP, ZMMSU). that such vicious creatures should bear my name.” In addition, two males from Repetek are housed in

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 11

TJK, Baljuvon TKM, Chainury Dimensions (mm) ♂ holotype (ZMUH) ♀ (FKCP) ♀ (FKCP) Carapace L / Wp 3.05 / 3.80 3.67 / 4.30 3.55 / 4.03 Mesosoma L 7.70 9.40 7.10 Tergite VII L / W 2.15 / 3.50 2.35 / 4.10 1.79 / 4.18 Metasoma + Telson L 23.38 23.41 21.48 Segment I L / W / D 2.80 / 1.61 / 1.43 2.80 / 1.95 / 1.79 2.50 / 1.75 / 1.43 Segment II L / W / D 3.40 / 1.60 / 1.58 3.48 / 1.78 / 1.85 3.00 / 1.63 / 1.58 Segment III L / W / D 3.80 / 1.73 / 1.73 3.85 / 1.85 / 1.90 3.43 / 1.68 / 1.73 Segment IV L / W / D 4.43 / 1.73 / 1.74 4.35 / 1.88 / 1.95 4.05 / 1.75 / 1.74 Segment V L / W / D 4.55 / 1.68 / 1.50 4.78 / 1.97 / 1.72 4.45 / 1.78 / 1.50 Telson L 4.40 4.15 4.05 Vesicle L / W / D 2.63 / 1.05 / 1.10 2.65 / 1.40 / 1.30 2.60 / 1.30 / 1.20 Aculeus L 1.77 1.50 1.45 Pedipalp L 11.08 11.15 10.65 Femur L / W 3.03 / 0.65 2.95 / 0.75 2.75 / 0.70 Patella L / W 3.40 / 0.95 3.40 / 1.08 3.25 / 1.00 Chela L 4.65 4.80 4.65 Manus L / W / D 1.35 / 0.75 / 0.70 1.35 / 0.86 / 0.85 1.11 / 0.75 / 0.75 Movable finger L 3.30 3.45 3.54 Total L 34.13 36.48 32.13

Table 1: Measurements of three adults of Anomalobuthus rickmersi. Abbreviations: length (L), width (W), posterior width (Wp), depth (D).

NHMW and one in R. Kinzelbach's collection (ZSRO). None of these crucial characters match the holotype Some of these specimens are illustrated here in Figs. 4– of A. rickmersi and clearly contradict the original des- 5, 27–30, 103–115. A few specimens from northern cription and illustrations published by Kraepelin (1900), Turkmenistan (Kunya-Urgench, ZISP 1699; Shakhs- which were confirmed by our own examination of the enem, ZMMSU Tb-374), published by Fet (1989) as A. true type: it is a large adult male with a total length of 34 rickmersi, actually correspond to a different species (A. mm, pectinal tooth count 25/25, fingers with 10 rows of pavlovskyi sp. n., see below). primary denticles of which the basal-most extends onto It is clear from A. Birula's publications that he never less than one-third of finger, and fulcra normally de- studied the holotype of A. rickmersi; however, it was veloped and pearl-like. Finally, a photo of the label of seen and its trichobothrial pattern sketched by Vachon the specimen examined by Lourenço (2001), provided (1974: 908, fig. 37). On the other hand, Lourenço (2001) by himself to V.F.in 2009, does not correspond to any of claimed to have examined it; however, it is obvious that those accompanying the actual holotype (Fig. 3). the specimen he received on loan from ZMUH was not Kraepelin's original holotype, but a misidentified and Anomalobuthus krivochatskyi Teruel, Kovařík et wrongly labeled juvenile of maybe even another genus. Fet, sp. n. Lourenço (2001) declared this specimen as an immature Figures 34–64, 120–125, 128–133, 134–136, 138–141. with a total length of 13–14 mm [sic], pectinal tooth Tables 2, 4–5 count 24–26, and textually wrote: “... cutting edge of the movable finger with 6 series of granules [...] the basal- http://zoobank.org/urn:lsid:zoobank.org:act:3936A6 most extending onto more than half the length of the 8B-96D5-461C-9568-5077BEA67488 finger. [... Pectinal] Fulcra almost vestigial and flat...” [originally in French, English translation and bracketed Anomalobuthus rickmersi: Soleglad & Fet, 2003a: 5, fig. text added by us]. 4 (in part; Bukhara); Soleglad & Fet, 2003b: 7, 59, 12 Euscorpius — 2018, No. 270

Figures 34–37: Anomalobuthus krivochatskyi sp. n., adult male holotype (34–35) and adult female paratopotype (36–37): full- body views, dorsal (34, 36) and ventral (35, 37). Scale bar = 10 mm.

156, tab. 2 (in part; Bukhara); Fet et al., 2005: 2, 6, dini, 2014: S1: 2, S2: 2; Teruel et al., 2014: 1, 4, 6–7, 9, 11; fig. 10 (in part; Bukhara); Fet et al., 2006: 2, 10, fig. 20, tabs. 1–2 (in part; Uzbekistan). tab. 3; Graham et al., 2012: 95–106, figs. 1–2, 6 (in HOLOTYPE ♂ (FKCP). Uzbekistan, Navoyi Pro- part; between Bukhara and Gazli); Loria & Pren- vince, Tamdy (=Tomdi) District, 1–2 km SSE of

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 13

Figures 38–41: Anomalobuthus krivochatskyi sp. n., adult male (38–39) and adult female paratypes (40–41) from between Bukhara and Gazli, Uzbekistan: full-body views, dorsal (38, 40) and ventral (39, 41). Scale bar = 10 mm.

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Zarafshan, 41º32'N 64º12'E, 18–28 April 1998, leg. A. with a diffuse annulated appearance (basal and distal V. Gromov. parts of each segment infuscate in the shape of thin blackish rings, which become somewhat larger and more PARATYPES. Same data as holotype, 1♂, 5♀♀ diffuse dorsally and basally), segment V entirely (FKCP), 1♂, 1♀ (RTOC: Sco-0233). Uzbekistan, Kizy- blackish; carinae not infuscate nor underlined with dark lkum Desert, Bukhara Province, between Bukhara and pigment, except for darkened terminal denticles on dor- Gazli, 40º05'N 64º04'E, 206 m a.s.l., 11 May 2002, leg. sal laterals and lateral supramedians. Telson vesicle V. Fet, 1♂, 1♀, 4 juveniles (NMPC), 1 juvenile blackish, similar to metasomal segment V; aculeus with (ZMUH), 1 juvenile (SOFM), 1♀ (RTOC: Sco-0208); basal half brownish and distal half blackish. Bukhara, 30 April 1979, leg. Antuš, 1♀ (FKCP). Chelicerae (Fig. 46). Manus smooth and shiny, Kazakhstan, South Kazakhstan Province, 5 km west of dorsodistal portion with some weak granules arranged Chardara, 41°16'14"N 67°53'02"E, 250 m a.s.l., 23–24 transversally, defining a depressed area; setation very May 2016, leg. P. Kučera, 1♀, 1 juvenile (FKCP). dense ventrally but essentially lacking dorsally, except for many rigid macrosetae on fixed finger and a few ETYMOLOGY. We are pleased to name this species around depressed area of manus. Fixed finger of size, after our colleague, Viktor Anatolyevich Krivochatsky shape and curvature ordinary for the family, dorsally (b. 1954) (St. Petersburg, Russia), an entomologist, a with the four standard denticles: single d very large, specialist in ant-lions (Neuroptera: Myrmeleontidae) and sharp and falcate, single sd large, sharp and falcate, an old friend of V.F. from the time of their work in the single m and b large, sharp, triangular and fused into a natural reserves of Turkmenistan in the 1970s-1980s. subrectangular bicusp; ventrally with a single denticle, Since 1991, Dr. Krivochatsky works in ZISP and is in which is medium-sized and placed at a level between charge of the famed scorpion collection of Alexei Birula. dorsal denticles m and b. Movable finger of size, shape and curvature ordinary for the family, dorsally with the DIAGNOSIS. Adult size 23–32 mm in males, 23–37 five standard denticles: single ed very large, sharp, mm in females. Coloration orange yellow, essentially falcate and placed at the same level of id, both closely immaculate, only with some regular blackish spots on opposable to fixed finger distal denticle, single sd large, pedipalps and metasoma (slightly darker and more sharp, triangular, single m very large, sharp, triangular, extended in female); metasomal segment V and telson two ed small (basalmost smaller), sharp, triangular and blackish. Pedipalp fingers with 9–11 principal rows of adjacent to each other and m; ventrally with a single denticles and 6–7 internal accessory denticles. Pectines denticle which is medium-sized, small, placed at the with 22–25 teeth in males and 19–22 in females. Tibial level of dorsal denticle sd and raises from a conspicuous, spur in males absent, in females present in legs III–IV. translucent lamella. Metasoma slightly attenuate, with most carinae weak; Pedipalps (Figs. 34–35). Relatively short but very ventral lateral carinae of segment V composed of slender, essentially bare. Femur subtly curved inwards, narrowly conical (males) to broadly lobate (females) with carinae weak, granulose to subdenticulate; denticles, dorsal lateral and lateral supramedian carinae intercarinal tegument smooth and glossy. Patella some- of segments I–IV with terminal denticle enlarged; what curved inwards, with carinae obsolete to vestigial, intercarinal areas essentially smooth, except sparsely smooth; intercarinal tegument smooth and glossy. Chela granulose on ventrodistal and ventrolateral areas of elongate and very slender; manus conspicuously nar- segment V. Telson vesicle oval and densely setose. rower than patella (ratio 0.78), cylindrical (2.14 times longer than wide, as wide as deep), with carinae obsolete DESCRIPTION (adult male holotype). Coloration to vestigial, smooth; intercarinal tegument smooth and (Figs. 34–35) base very vivid yellow, with an orange glossy; fingers very long (movable finger 2.20 times shade; in general, the base color is paler on pedipalp longer than underhand), only subtly curved and with 9– chelae, legs, and pectines. Chelicerae immaculate, ex- 10 principal rows of denticles (the two basalmost rows cept for blackish finger teeth. Pedipalp femur with a are poorly defined), basal lobe/notch combination ab- conspicuous blackish spot on dorsodistal apex, which sent, external accessory denticles absent, internal acces- diffusely continues as a broad stripe over almost all sory denticles very large and claw-like (increasing in internal surface; patella with internal surface blackish; size distally), numbering seven and six on fixed and chela immaculate, only with finger denticles blackish. movable fingers, respectively, movable finger with one Carapace immaculate, only with a blackish spot under claw-like accessory denticles basal to the very large every ocular group. Tergites immaculate. Coxosternal terminal denticle. region and genital operculum immaculate. Pectines pale Carapace (Fig. 46). Very strongly trapezoidal yellowish, immaculate. Sternites immaculate. Legs im- (much narrower anteriorly) and wider than long; anterior maculate; claws with distal half dark brown to blackish. margin shallowly convex, with 6–7 pairs of thin Metasoma conspicuously bicolor: segments I–IV yellow macrosetae and some very short microsetae. Carination

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 15

Figures 42–45: Anomalobuthus krivochatskyi sp. n., adult female (42–43) and juvenile male paratypes (44–45) from Chardara, Kazakhstan: full-body views, dorsal (42, 44) and ventral (43, 45). essentially absent: the only carinae present are the trals, central transverse, and posterior laterals long, superciliaries, which are weakly granulose to smooth. narrow and shallow. Tegument very finely and densely Furrows: anterior median, median ocular, central me- granulose, with many small to medium-sized granules dian, posterior median and posterior marginal fused, scattered all over except on both sides of median axis, wide and moderately deep; lateral oculars, lateral cen- where the granules are coarser and glossy.

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Figures 46–53: Anomalobuthus krivochatskyi sp. n., adult male holotype (46–47) and adult female paratopotype (48–53): chelicerae, carapace and tergites I–III, dorsal (46, 48), sternopectinal region and sternites III–V, ventral (47, 49), distal part of right legs I–IV, internal (50–53).

Sternum (Fig. 47). Standard for the genus, re- teromedian furrow, posterior margin very shallowly latively small and widely triangular in shape, with two convex. pairs of inconspicuous macrosetae. Legs (Figs. 34–35). Very slender, with all carinae Genital operculum (Fig. 47). Relatively large, each weak to obsolete and subgranulose to smooth; inter- half roundly subtriangular in shape, with three pairs of carinal tegument smooth and glossy. Tibial spur absent. inconspicuous macrosetae, plus a few short microsetae. Mesosoma (Figs. 46–47). Tergites with the same Genital papillae present. sculpture as on carapace; I–VI irregularly tricarinate: the Pectines (Fig. 47). Standard-sized for the genus: median longitudinal carina is moderately strong, short, very long, extending beyond leg IV coxa-trochanter and formed by irregular medium-sized granules that do joint), subrectangular and densely setose. Tooth count not project beyond posterior margin, but the submedian 22/22. Basal plate heavily sclerotized, much wider than carinae are undefined on I–IV, very irregular on V, and long, anterior margin with a very deep, narrow an- well defined only on VI; tergite VII with five well-

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 17

Figures 54–58: Anomalobuthus krivochatskyi sp. n., adult female paratopotype: right pedipalp trochanter and femur, dorsal (54), right pedipalp patella, dorsal (55) and external (56), right pedipalp chela, dorsoexternal (57), right pedipalp movable finger, dorsoexternal (58). Black dots depict trichobothria. defined carinae (median, submedians and laterals), denticulate granulation: dorsal laterals obsolete to ves- which are long, strong and finely serrate to crenulate. tigial on I–IV (with one terminal denticles enlarged), Sternites essentially bare; III–VI glossy and with subtle absent on V; lateral supramedians obsolete to vestigial vestiges of a pair of smooth submedian carinae, spiracles on I–IV (with 1–2 terminal denticles enlarged), vestigial relatively short and slit-like, transversely arranged (not as rounded ridges on V; lateral inframedians obsolete to oblique), V with smooth patch absent; VII with two vestigial on I–III, absent on IV–V; ventral laterals pairs of carinae (submedians and laterals) which are long obsolete to vestigial on I–IV, moderate to strong on V, and finely crenulate to serrate, intercarinal tegument where become progressively stronger and somewhat coriaceous to minutely granulose. flared distally, formed by sharp, subequal denticles; Metasoma (Figs. 59–61). Somewhat elongated and ventral submedians moderate on I–II, weak on III, slightly wider both basally and distally; with 10/10/ obsolete on IV, absent on V (indicated by somewhat 10/8/5 complete to almost complete carinae, almost all raised tegument and irregular granulation on basal half); formed by conspicuously isolated, sharply serrate to ventral median absent on I–IV, moderate on V. Inter- 18 Euscorpius — 2018, No. 270

Figures 59–64: Anomalobuthus krivochatskyi sp. n., adult male holotype (59–61) and adult female paratopotype (62–64): metasoma and telson, lateral (59, 62), dorsal (60, 63) and ventral (61, 64). carinal tegument smooth and glossy, with sparse with vestiges of coarse granules arranged into three granulation of different sizes ventrally. Dorsal furrow obsolete longitudinal carinae (ventral median and ventral wide and shallow on all segments. Setation sparse, submedians), and some coarse punctations ventrally. mostly represented by 4–7 dark macrosetae over every Subaculear tubercle absent, but subtly suggested by a carina. vestigial granule. Aculeus conspicuously shorter than Telson (Figs. 59–61). Moderately setose, with many vesicle, thick and shallowly curved. setae scattered all over dorsal and lateral surfaces. Vesicle short oval (1.92 times longer than wide, 0.92 FEMALE (paratopotype). Very similar to described times wider than deep), tegument smooth and glossy, male, sexual dimorphism evident by: 1) size compara-

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 19

UZB, Bukhara to UZB, Zarafshan Gazli Dimensions (mm) ♂ holotype ♂ paratype ♀ paratype ♀ paratype (FKCP) (RTO: Sco-0233) (RTO: Sco-0233) (RTO: Sco-0208) Carapace L / Wp 3.30 / 3.70 3.40 / 3.80 3.80 / 4.60 3.00 / 3.40 Mesosoma L 7.00 6.80 9.50 7.00 Tergite VII L / W 2.22 / 3.40 2.00 / 3.60 2.20 / 4.60 1.80 / 3.70 Metasoma + Telson L 21.90 21.30 23.30 18.40 Segment I L / W / D 2.60 / 1.93 / 1.60 2.40 / 1.90 / 1.62 2.50 / 2.30 / 1.96 2.10 / 1.70 / 1.47 Segment II L / W / D 3.13 / 1.78 / 1.55 3.00 / 1.70 / 1.70 3.30 / 2.20 / 2.00 2.60 / 1.60 / 1.48 Segment III L / W / D 3.45 / 1.85 / 1.65 3.30 / 1.80 / 1.77 3.60 / 2.20 / 2.07 2.90 / 1.70 / 1.50 Segment IV L / W / D 4.04 / 1.89 / 1.60 4.00 / 1.90 / 1.75 4.40 / 2.40 / 2.11 3.40 / 1.80 / 1.51 Segment V L / W / D 4.48 / 1.92 / 1.42 4.40 / 1.90 / 1.55 4.80 / 2.40 / 1.80 3.80 / 1.80 / 1.41 Telson L 4.20 4.20 4.70 3.60 Vesicle L / W / D 2.75 / 1.28 / 1.26 2.30 / 1.20 / 1.30 2.70 / 1.50 / 1.40 2.10 / 1.20 / 1.10 Aculeus L 1.45 1.90 2.00 1.50 Pedipalp L 10.51 11.00 11.40 9.50 Femur L / W 2.78 / 0.68 2.80 / 0.70 3.10 / 0.90 2.50 / 0.70 Patella L / W 3.33 / 0.93 3.40 / 0.90 3.60 / 1.10 2.90 / 0.90 Chela L 4.40 4.80 4.70 4.10 Manus L / W / D 1.22 / 0.71 / 0.75 1.50 / 0.70 / 0.70 1.40 / 0.80 / 0.90 1.20 / 0.70 / 0.70 Movable finger L 3.18 3.30 3.30 2.90 Total L 32.20 31.50 36.60 28.40

Table 2: Measurements of four types of Anomalobuthus krivochatskyi sp. n. Abbreviations: length (L), width (W), posterior width (Wp), depth (D).

tively larger inside each size-class; 2) mesosoma and Specimens from Chardara and Bukhara to Gazli are metasoma slightly less slender; 3) genital papillae ab- identical to those from the type locality in all diagnostic sent; 4) pectines smaller, with consistently lower tooth characters. They only differ by being slightly smaller counts; 5) tibial spur small, sharp and sinuose, present at and darker, with better-marked dark patterns (Figs. 38– least on both legs IV (sometimes also on one leg III); 6) 45). These differences are minor and attributable to metasomal segment V with ventral intercarinal tegument standard variation between different populations. less granulose. See Figs. 36–37, 40–43, 48–58, 62–64, 134–136, 138–140 and Tabs. 2, 4–5. COMPARISON. Adults of A. krivochatskyi sp. n. can be very easily distinguished by the almost absolute VARIATION. Adult size varied from 31.5–32.2 mm smoothness of the metasomal intercarinal spaces, es- in males and 28.4–36.6 mm in females. pecially in males (Figs. 59–64, 121); almost all other Count of teeth per pecten varied as follows (Tab. 5): congeners have at least the ventral and lateral surfaces of in males 22 (3), 23 (1), 24 (3), 25 (3), and in females 19 segments IV–V moderately to densely granulose (Figs. (2), 20 (7), 21 (6), 22 (3). No significant differences de- 23–25, 27–29, 31–33, 69, 75–78, 84–89). tected among specimens from the localities examined by The single exception is A. talebii, which is similarly us. smooth (Figs. 98–100), but the only female known has The tibial spur is always absent in males, but in much higher pectinal tooth count (26/26 vs. 20–23) and females it is present always on both leg IV and occasion- several quite distinct morphometric ratios (see Tab. 4 ally also in leg III of one side only (Figs. 130–132, 134). and Comparison section of that species).

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DISTRIBUTION (Fig. 137). Sands of the Kizylkum non-optimal preservation. Chelicerae immaculate, ex- Desert, from central Uzbekistan through adjacent sou- cept for blackish finger teeth. Pedipalp femur with a thern edge of Kazakhstan. conspicuous blackish spot on dorsodistal apex, which diffusely continues as a broad stripe over almost all NOTES. The Bukhara Province (between Bukhara internal surface; patella with internal surface blackish; and Gazli) specimens of this new species were included chela with internal surface diffusely reticulate with dark (under A. rickmersi) in a DNA phylogeographic study by brown, finger denticles blackish. Carapace with a large Graham et al. (2012), which demonstrated a derived blackish spot that widely surrounds median ocular position of this taxon within the genus. tubercle and extends to anterior margin and also dif- fusely to every group of lateral eyes. Tergites diffusely Anomalobuthus lowei Teruel, Kovařík et Fet, sp. n. marbled with dark brown. Coxosternal region and Figures 65–78, 141; Tables 3–5 genital operculum immaculate. Pectines pale yellowish, immaculate. Sternites immaculate. Legs spotted with http://zoobank.org/urn:lsid:zoobank.org:act:F416B1 dark brown, more densely on trochanter and femur of I– 5C-4E36-4D73-ABCC-DCCE1A6E9DB9 III; claws with distal third dark brown to blackish. Metasoma bicolor, segments I–IV yellow with a diffuse Anomalobuthus rickmersi: Gromov & Kopdykbaev, annulated appearance (basal and distal parts of each 1994: 20 (in part); Gromov, 2005: 39 (in part); segment infuscate in the shape of thin blackish rings, Gromov & Kazenas, 2006: 39 (in part); Mityaev et which become somewhat larger and more diffuse dor- al., 2005: 39 (in part); Gromov, 2006: 44–45 (in sally and basally) and segment V very densely reticu- part); Kamenz & Prendini, 2008: 6, 8, 40, 56–57, lated with blackish brown, becoming gradually paler and fig. 15a, tabs. 1–2. sparser towards basal half; carinae not infuscate nor underlined with dark pigment. Telson vesicle blackish, HOLOTYPE ♀ (GLPC). Kazakhstan, Almaty Prov- aculeus with basal half brownish and distal half blackish. ince, Kapchagay [43°53'N, 77°05'E], Ili River, 75 km Chelicerae (Fig. 67). With dentition typical for the north of Almaty, 17 May 1993, leg. A. Feodorov. genus, as described for A. krivochatskyi sp. n. (see above). ETYMOLOGY. We are pleased to name this species Pedipalps (Figs. 70–74). Relatively short but after our friend and colleague, Graeme Lowe (Phil- slender, essentially bare. Trichobothrial pattern A-β neo- adelphia, USA), who contributed greatly to the bothriotaxic, with the following trichobothria missing: knowledge of the Palearctic desert scorpions. His assis- femoral d2 and chelal esb (both pedipalps), chelal Et and tance to the present paper was crucial, by selflessly chelal V1 (right pedipalp). Chelal it in subdistal position, spending his scarce free time in exhausting photographic midway between second and third enlarged principal sessions providing all the images of the holotype of A. denticle from tip (left chela) or just distal to third lowei (Figs. 65–78). enlarged principal denticle from tip (right chela). Femur essentially straight, with carinae weak to moderate, gran- DIAGNOSIS (based on one adult female). Adult size ulose to subdenticulate; intercarinal tegument smooth 35 mm. Coloration yellow, essentially immaculate, only and glossy. Patella essentially straight, with carinae with some irregular blackish spots on carapace, tergites, obsolete to absent, smooth; intercarinal tegument smooth pedipalps, legs, and metasoma; metasomal segment V and glossy. Chela elongate and slender; manus con- deeply infuscate, telson blackish. Pedipalp fingers with spicuously narrower than patella (ratio 0.70), cylindrical 7–8 principal rows of denticles and 6–7 internal (1.94 times longer than wide, 1.13 times deeper than accessory denticles. Pectines with 21/20 teeth. Tibial wide), with carinae obsolete to absent, smooth; inter- spur on leg IV only, small. Metasoma robust, with most carinal tegument smooth and glossy; fingers long carinae moderately developed; ventral lateral carinae of (movable finger 2.31 times longer than underhand), only segment V composed of narrowly lobate denticles, subtly curved and with 8–9 principal rows of denticles dorsal lateral and lateral supramedian carinae of seg- (the two basalmost rows are poorly defined), basal ments I–IV with terminal denticle greatly enlarged; lobe/notch combination absent, external accessory den- intercarinal spaces essentially smooth on dorsal areas of ticles absent, internal accessory denticles very large and all segments, densely granulose on ventral and lateral claw-like (increasing in size distally), numbering seven areas of I and V, sparsely granulose on ventral and and six on fixed and movable fingers, respectively, lateral areas of II–IV. Telson vesicle short oval and movable finger with two claw-like accessory denticles sparsely setose. basal to the very large terminal denticle. Carapace (Fig. 67). Very strongly trapezoidal DESCRIPTION (adult female holotype). Coloration (much narrower anteriorly) and wider than long; anterior (Fig. 65) base light yellow, somewhat translucent due to margin shallowly convex, with 5–6 pairs of thin macro-

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 21

Figures 65–74: Anomalobuthus lowei sp. n., adult female holotype. Figures 65–66. full-body dorsal views, under white light (65) and under UV light (66). Figures 67–68. Mesosoma dorsal (67) and ventral (68) under UV light. Figure 69. Metasoma and telson under UV light, lateral. Figures 70–74. Pedipalp under UV light: right pedipalp trochanter and femur, dorsal (70), right pedipalp patella, dorsal (71), right pedipalp chela, dorsal (72), external (73) and ventral (74). Scale bars = 10 mm (65–66) and 1 mm (67–68, 70–74). Photographs courtesy Graeme Lowe. setae and some very short microsetae. Carination essen- rows: anterior median, median ocular, central median, tially absent: the only carinae present are the super- posterior median and posterior marginal fused, wide and ciliaries, which are vestigially granulose to smooth. Fur- very shallow; lateral oculars, lateral centrals, central

22 Euscorpius — 2018, No. 270

Figures 75–78: Anomalobuthus lowei sp. n., adult female holotype. Figures 75–76. Metasoma and telson, dorsal, under white light (75) and under UV light (76). Figures 77–78. Metasoma and telson, ventral, under white light (77) and under UV light (78). Scale bar = 2 mm. Photographs courtesy Graeme Lowe. transverse, and posterior laterals long, narrow and shal- from both legs III and right leg IV, but moderate-sized low. Tegument very finely and densely granulose, with on left leg IV. many small to medium-sized granules scattered all over Mesosoma (Figs. 67–68). Tergites with the same except on ocular triangle, where the granules are coarser sculpture as on carapace; I–VI essentially monocarinate: and glossy. the median longitudinal carina is moderately strong, Sternum (Fig. 68). Standard for the genus 1, very long, and formed by irregular medium-sized granules small, and narrowly pentagonal in shape, with two pairs that do not project beyond posterior margin and the of macrosetae. submedian carinae are undefined on I–VI, although Genital operculum (Fig. 68). Very large, each half suggested by a few slightly coarser, irregular granules; widely crescent-shaped, with 3–4 pairs of inconspicuous tergite VII with five well-defined carinae (median, sub- macrosetae, plus a few short microsetae. Genital papillae medians and laterals), which are long, strong and serrate absent. to denticulate. Sternites essentially bare except on Pectines (Fig. 68). Somewhat small for the genus posterior margin, which is conspicuously setose; III–VI (just reaching leg IV coxa-trochanter joint), subrec- acarinate, glossy but with posterior and lateral margins tangular and densely setose. Tooth count 21/20 (terminal finely and densely granulose, spiracles relatively short tooth of right pecten missing, but base remains allows and slit-like, transversely arranged (not oblique), V with unequivocal count). Basal plate heavily sclerotized and smooth patch absent; VII with two pairs of carinae: the densely granulose, much wider than long, anterior mar- submedians are long and finely denticulate and the gin with a very deep, narrow anteromedian furrow, laterals are short and coarsely denticulate, intercarinal posterior margin very shallowly convex. tegument very densely granulose. Legs (Figs. 65–66). Very slender, with all carinae Metasoma (Figs. 69, 75–78). Robust and slightly weak to obsolete and subgranulose to smooth; inter- wider both basally and distally; with 10/10/10/8/5 carinal tegument smooth and glossy. Tibial spurs absent complete to almost complete carinae, all except dorsal

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 23

laterals and laterals supramedians, formed by con- DISTRIBUTION (Fig. 141). Sands of the Ili River tiguous, sharply serrate to denticulate granulation: dorsal valley in southeastern Kazakhstan. It is the easternmost laterals moderate on I, strong on II–III, moderate on IV species of Anomalobuthus, occurring at the northern (with terminal denticle greatly enlarged as a coarse, piedmont of the Tian Shan Range. conical tubercle), absent on V; lateral supramedians moderately strong on I, weak on II–III, obsolete to very NOTES. Based on geographical occurrence, the male weak on IV (with 1–2 terminal denticles enlarged), recorded by Kamenz & Prendini (2008) as A. rickmersi vestigial as rounded ridges on V; lateral inframedians most likely corresponds to A. lowei sp. n. strong on I, moderate on II (undefined on basal 1/4), weak on III (undefined on basal 1/3), absent on IV–V; Anomalobuthus pavlovskyi Teruel, Kovařík et ventral laterals strong on I–II, moderate on III–IV, very Fet, sp. n. strong on V, where become progressively stronger and Figures 79–89, 107–116, 141; Tables 3–5 somewhat flared distally, formed by round, subequal denticles; ventral submedians very strong on I–III, http://zoobank.org/urn:lsid:zoobank.org:act:D8 strong on III, moderate on IV, absent on V; ventral B0A76D-BF70-426C-B822-D9E9C00A96C5 median absent on I–IV, moderate on V (poorly defined from intercarinal granulation). Intercarinal tegument Anomalobuthus rickmersi: Birula, 1911b: 171 (in smooth and glossy, with fine granules scattered all over part; “Bajgakum” [=Baigakum]); Pavlovsky, 1916a: 35 except dorsally on I–IV, on V such granules are re- (in part; Dzhulek); Pavlovsky, 1916b: 243, fig. 1 (in stricted to ventral surface but are much denser. Dorsal part; Dzhulek); Fet, 1989: 80, unnumbered figure in furrow wide and moderately deep on all segments. page 138 (in part; Dzhulek and Kunya-Urgench); Gro- Setation moderately dense, mostly represented by 6–8 mov & Kopdykbaev, 1994: 20 (in part; Dzhulek); dark macrosetae over every carina. Setae on lateral Gromov, 2005: 39 (in part); Mityaev et al., 2005: 39 (in surfaces of IV and V, and ventral surface of V arising part); Gromov, 2006: 44–45 (in part); Graham et al., from anterior edges of shallow pits or depressions. 2012: 95–106, figs. 1–2, 6 (in part; Baigakum). Lateral anal margins of V with 3 lobes, ventral anal arc heavily granulated. HOLOTYPE ♂ (RTOC: Sco-0476). Kazakhstan, Telson (Figs. 69, 75–78). Moderately setose, with Kizylkum Desert, Kyzyl-Orda Province, Chiili District, some setae scattered over dorsal and lateral surfaces. ca 2.5 km NW of Baigakum (=Djulek or Dzhulek), Vesicle short oval (1.47 times longer than wide, 1.13 44°20'37'' to 44°20'29''N, 66°27'09'' to 66°27'07''E, 127– times wider than deep), tegument smooth and glossy, 143 m a.s.l., 25 May 2002, leg. V. Fet & A. V. Gromov. with vestiges of coarse granules arranged into five vestigial longitudinal carinae (ventral median, ventral PARATYPES. Same data as holotype, 2♂♂, 2♀♀ submedians, and ventral laterals), and some coarse (FKCP), 1♂ (ZMUH), 1♂ (SOFM), 1♂ (NMPC), 3♀♀ punctations ventrally. Subaculear tubercle absent. (RTOC: Sco-0476). Same data as holotype except 6–21 Aculeus shorter than vesicle, thick and moderately June [19 June–4 July] 1907, leg. D. Glazunov, 2♂♂ curved. (ZISP 810, one examined by Max Vachon and bearing his handwritten code VA-1667). Turkmenistan, Daşo- MALE. Unknown. guz (=Tashauz), Province, Köneürgenç (=Kunya- Urgench), 42º23'18"N 59º16'14"E, 3 June 1972, leg. G. VARIATION. Unknown, no other specimens avail- S. Medvedev, 1♂, 1♀ (ZISP 1699). able to us. OTHER MATERIAL. Turkmenistan, Daşoguz (=Tas- COMPARISON (females only). A. lowei sp. n. can be hauz) Province, Shakhsenem [old fortress ruins], 8 very easily distinguished from all other Anomalobuthus October 1983, leg. O. Soyunov, 1♀ (ZMMSU Tb-374). spp. by its very robust appearance, well evident in its many unique morphometric proportions (Tab. 4). For ETYMOLOGY. We dedicate this species to the mem- example, it is the only species of this genus with ory of the famous Russian zoologist Evgenii Nik- metasomal segment I about as long as wide (ratio = 1.02 anorovich Pavlovsky (1884–1965), who produced many vs. 1.09–1.44 in all other congeners), it has the shortest well-known contributions to scorpion anatomy and metasoma + telson when compared to carapace length histology. He was also the Director of ZISP for 20 years (ratio = 5.40 vs. 5.68–6.38), the shortest metasomal seg- (after Alexei Birula was arrested in 1931 during the ment V (length/depth ratio = 1.85 vs. 2.00–2.50) and the purges), and was the first one to observe and collect this deepest telson vesicle (length/depth ratio = 1.66 vs. species at the type locality, more than 100 years ago 1.89–2.17). (Pavlovsky, 1916a, 1916b). 24 Euscorpius — 2018, No. 270

Figures 79–82: Anomalobuthus pavlovskyi sp. n., adult male (79–80) and adult female paratopotypes (81–82): full-body views, dorsal (79, 81) and ventral (80, 82). Scale bar = 10 mm.

DIAGNOSIS. Adult size 22–30 mm in males, 23–36 female); metasomal segment V bicolor, blackish with mm in females. Coloration light yellow, essentially im- yellowish red basal third (male) to basal half (female), maculate, only with some regular blackish spots on telson blackish. Pedipalp fingers with 8–9 principal rows pedipalps and metasoma (darker and more extended in of denticles and 7–8 internal accessory denticles. Pec-

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 25 tines with 24–27 teeth in males and 22–23 in females. internal accessory denticles very large and claw-like Tibial spur in males entirely absent to present only in leg (increasing in size distally), numbering eight and seven IV, in females absent to present in legs III–IV. Meta- on fixed and movable fingers, respectively, movable soma moderately robust, with most carinae obsolete to finger with three claw-like accessory denticles basal to vestigial; ventral lateral carinae of segment V composed the very large terminal denticle. of narrowly conical denticles, dorsal lateral and lateral Carapace. Very strongly trapezoidal (much nar- supramedian carinae of segments I–IV with terminal rower anteriorly) and wider than long; anterior margin denticle enlarged; intercarinal areas of segments I–IV straight, with 5–6 pairs of thin macrosetae and some with granulation sparse ventrally (male) to dense very short microsetae. Carination essentially absent: the ventrally and laterally (female), of segment V moder- only carinae present are the superciliaries, which are ately dense ventrally (male) to very dense ventrally and weakly granulose. Furrows: anterior median, median laterally (female). Telson vesicle elongate oval and ocular, central median, posterior median and posterior sparsely setose. marginal fused, wide and moderately deep; lateral ocu- lars, lateral centrals, central transverse, and posterior DESCRIPTION (adult male holotype, except figures laterals long, narrow and moderately deep. Tegument of adult male paratopotype). Coloration (Figs. 79–80) very finely and densely granulose, with several small to base light yellowish, with a subtle orange shade; in medium-sized granules scattered all over general, the base color is paler on pedipalp chelae, legs, Sternum. Standard for the genus, relatively small and pectines. Chelicerae subtly reticulate with dark and widely triangular in shape, with one pair of in- brown, finger teeth blackish. Pedipalp femur with a conspicuous macrosetae. conspicuous brown spot on dorsodistal apex, which Genital operculum. Relatively large, each half diffusely continues as a broad stripe over almost all roundly subtriangular in shape, without macrosetae but internal surface; patella with internal surface brown; with a few short microsetae. Genital papillae present. chela immaculate, only with finger denticles blackish. Pectines. Standard-sized for the genus: long, Carapace largely immaculate, only with a blackish spot extending beyond leg IV coxa-trochanter joint), sub- under every ocular group and interocular triangle mod- rectangular and densely setose. Tooth count 26/27. Basal erately infuscate. Tergites immaculate. Coxosternal re- plate heavily sclerotized, much wider than long, anterior gion and genital operculum immaculate. Pectines pale margin with a very deep, narrow anteromedian furrow, yellowish, immaculate. Sternites immaculate. Legs im- posterior margin very shallowly convex. maculate; claws with distal half dark brown. Metasoma Legs (Figs. 79–80). Very slender, with all carinae bicolor, segment V blackish with basal third yellowish weak to obsolete and subgranulose to smooth; inter- red, I–IV yellow with a diffuse annulated appearance: carinal tegument smooth and glossy. Tibial spurs present basal and distal parts of each segment infuscate in the only on right leg IV. shape of thin blackish rings, which become somewhat Mesosoma (Figs. 79–80). Tergites with the same larger and more diffuse dorsally and basally; carinae not sculpture as on carapace; I–VI irregularly tricarinate: the infuscate nor underlined with dark pigment, except for median longitudinal carina is moderately strong, short, darkened terminal denticles on dorsal laterals and lateral and formed by irregular medium-sized granules that do supramedians. Telson blackish, with basal half of not project beyond posterior margin, but the submedian aculeus yellowish. carinae are poorly defined on I–VI; tergite VII with five Chelicerae. With dentition typical for the genus, as well-defined carinae (median, submedians and laterals), described for A. krivochatskyi sp. n. (see above). which are long, strong and finely serrate. Sternites Pedipalps (Fig. 83). Relatively short but very slen- essentially bare; III–VI glossy and with a pair of weak, der, essentially bare. Femur subtly curved inwards, with smooth submedian carinae, spiracles relatively short and carinae weak, granulose to subdenticulate; intercarinal slit-like, transversely arranged (not oblique), V with tegument smooth and glossy. Patella straight, with smooth patch absent; VII with two pairs of carinae carinae vestigial to weak, smooth; intercarinal tegument (submedians and laterals) which are long and finely smooth and glossy. Chela elongate and very slender; crenulate, intercarinal tegument coriaceous to minutely manus conspicuously narrower than patella (ratio 0.71), granulose. cylindrical (2.00 times longer than wide, 1.20 times Metasoma (Figs. 84–86). Slightly elongated and deeper than wide), with carinae vestigial to weak, essentially parallel-sided; with 10/10/10/8/5 complete to smooth; intercarinal tegument smooth and glossy; fin- almost complete carinae, most formed by conspicuously gers very long (movable finger 2.40 times longer than isolated, sharply serrate to denticulate granulation: dor- underhand), only subtly curved and with 8/9 principal sal laterals vestigial on I–III, obsolete on IV (with 1–2 rows of denticles (the two basalmost rows are poorly terminal denticle slightly enlarged), absent on V; lateral defined on movable finger), basal lobe/notch com- supramedians vestigial to obsolete on I–IV (with 1–2 bination absent, external accessory denticles absent, terminal denticles slightly enlarged), vestigial as round- 26 Euscorpius — 2018, No. 270

Figures 83–89: Anomalobuthus pavlovskyi sp. n., adult male (83–86) and adult female paratopotypes (87–89): right pedipalp (83), metasoma and telson, lateral (84, 87), dorsal (85, 88) and ventral (86, 89). ed ridges on V; lateral inframedians weak on I, obsolete formed by sharp, subequal denticles; ventral submedians to vestigial on II–III, absent on IV–V; ventral laterals weak on I–III, vestigial on IV, absent on V (indicated by weak on I–IV, moderate to strong on V, where become somewhat raised tegument and irregular granulation on progressively stronger and somewhat flared distally, basal half, but largely obscured by intercarinal gran-

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 27

A. lowei sp. n. A. pavlovskyi sp. n. (RTO: Sco-0476) DIMENSIONS (MM) ♀ holotype ♂ holotype ♀ paratype ♀ paratype Carapace L / Wp 3.88 / 4.75 2.10 / 2.90 2.60 / 3.00 3.88 / 4.47 Mesosoma L 10.03 5.50 5.60 9.68 Tergite VII L / W 2.41 / 4.79 1.60 / 2.08 1.50 / 2.80 2.30 / 4.28 Metasoma + Telson L 20.97 15.00 15.00 22.02 Segment I L / W / D 2.41 / 2.37 / 1.92 1.70 / 1.30 / 1.13 1.70 / 1.30 / 1.17 2.41 / 1.98 / 1.68 Segment II L / W / D 3.02 / 2.23 / 1.95 2.10 / 1.20 / 1.18 2.20 / 1.20 / 1.19 3.13 / 1.80 / 1.83 Segment III L / W / D 3.25 / 2.33 / 2.06 2.30 / 1.30 / 1.20 2.40 / 1.30 / 1.20 3.48 / 1.87 / 1.88 Segment IV L / W / D 4.08 / 2.33 / 2.14 2.90 / 1.30 / 1.21 2.80 / 1.30 / 1.28 4.10 / 2.00 / 2.00 Segment V L / W / D 4.33 / 2.34 / 1.84 3.10 / 1.30 / 1.19 3.00 / 1.30 / 1.27 4.50 / 2.08 / 1.73 Telson L 4.08 2.90 2.90 4.40 Vesicle L / W / D 2.33 / 1.58 / 1.40 1.70 / 0.90 / 0.80 1.70 / 0.90 / 0.80 2.50 / 1.51 / 1.20 Aculeus L 1.67 1.20 1.20 1.90 Pedipalp L 12.01 7.90 8.00 11.44 Femur L / W 3.17 / 0.92 2.00 / 0.50 2.00 / 0.60 2.90 / 0.81 Patella L / W 3.58 / 1.17 2.50 / 0.70 2.50 / 0.80 3.50 / 1.10 Chela L 5.26 3.40 3.50 5.04 Manus L / W / D 1.59 / 0.82 / 0.93 1.00 / 0.50 / 0.60 1.00 / 0.60 / 0.60 1.42 / 0.73 / 0.90 Movable finger L 3.67 2.40 2.50 3.62 Total L 34.88 22.60 23.20 35.58

Table 3: Measurements of the types of two new species of Anomalobuthus. Abbreviations: length (L), width (W), posterior width (Wp), depth (D). ulation); ventral median absent on I–IV, moderate on V and metasoma slightly less slender; 3) genital papillae but largely obscured by intercarinal granulation. Inter- absent; 4) pectines smaller, with consistently lower tooth carinal tegument smooth and glossy, with sparse counts; 5) metasomal segments with ventral and lateral granulation of different sizes ventrally. Dorsal furrow intercarinal tegument much more densely granulose. See wide and shallow on all segments. Setation moderately Figs. 81–82, 87–89, 114 and Tabs. 3–5 dense, represented by 6–9 mostly pale macrosetae over every carina. VARIATION. Adult size varied from 22.6–25.5 mm Telson (Figs. 84–86). Sparsely setose, with some in males and 23.2–35.6 mm in females. setae scattered all over dorsal and lateral surfaces. Pectinal tooth count varied as follows (Tab. 5): in Vesicle short oval (2.00 times longer than wide, 0.83 males 25 (5), 26 (5), 27 (2), and in females 22 (5) and 23 times wider than deep), tegument smooth and glossy, (3). No significant differences detected among speci- with vestiges of coarse granules arranged into three mens from the localities examined by us. obsolete longitudinal carinae (ventral median and ventral The tibial spur is highly variable amongst paratypes: submedians), some also on dorsal surface, and some it is entirely absent from all legs (one male), present on a coarse punctations all over. Subaculear tubercle absent, single leg IV (three males including holotype, two but subtly suggested by a vestigial granule. Aculeus females), present on each leg III–IV of the same side conspicuously shorter than vesicle, thick and shallowly (left, one female), or present on both legs IV (two males, curved. one female). In all cases where it is present, it is small to vestigial. FEMALE (paratopotype). Very similar to described male, sexual dimorphism evident by: 1) size com- COMPARISON. A. pavlovskyi sp. n. can be very paratively larger inside each size-class; 2) mesosoma easily distinguished by having intercarinal spaces of 28 Euscorpius — 2018, No. 270 metasoma much more densely granulose in females than stripes; metasomal segment V and telson blackish. in males (Figs. 84–89); in all other Anomalobuthus spp., Pedipalp fingers with 10–11 principal rows of denticles the metasomal segments are mostly sparsely granulated and 6–7 internal accessory denticles. Pectines with 26/26 to smooth and glossy in females (A. krivochatskyi sp. n., teeth. Tibial spur absent. Metasoma slightly attenuate, A. lowei sp. n. and A. talebii, see Figs. 62–64, 69, 75– with most carinae moderately developed; ventral lateral 78, 98–100), or more densely granulose in males (A. carinae of segment V composed of narrowly conical rickmersi, see Figs. 23–25, 27–29, 31–33). denticles, dorsal lateral and lateral supramedian carinae Moreover, this species and A. lowei sp. n. are the of segments I–IV with terminal denticle greatly en- only that still retain as adult a typically juvenile char- larged; intercarinal spaces essentially smooth on all acter: the segment V with basal third yellowish red segments, except for very few sparse granules on ven- (Figs. 84–89); in all other congeners it is entirely reddish trodistal area of V. Telson vesicle short oval and or blackish, or only subtly faded to dark reddish brown sparsely setose. at it extreme base (Figs. 1–2, 4–5, 8–9, 23–45, 59–64, 90–91, 98–100). But A. lowei sp. n. is very different in DESCRIPTION. See Teruel et al. (2014). many morphometric ratios (see Tab. 4 and Comparison section of that species). MALE. Unknown.

DISTRIBUTION (Fig. 141). Lowland sands of the VARIATION. Unknown, single specimen known. Aral Basin (Kizylkum Desert), from south-central Kaz- akhstan through extreme northern Turkmenistan. COMPARISON (females only). A. talebii can be unequivocally distinguished by its much higher pectinal NOTES. tooth count of 26/26, while all other Anomalobuthus spp. do have 19–23 teeth (Tab. 5). Also, it has some unique 1. The first specimens of this species were recorded by morphometric proportions, e.g., the slenderest telson Birula (1911b) as A. rickmersi; all were collected from vesicle (length/width ratio = 2.18 vs. 1.47–2.00 in the Baigakum Sands (Dzhulek) in 1907 by Dmitri Kons- other species) and the shortest pedipalp movable finger tantinovich Glazunov (1869–1914), a Russian ento- when compared to manus length (ratio = 2.26 vs. 2.31– mologist and brother of the composer Alexander K. 3.18 in all other congeners); see Tab. 4. Glazunov. It is the only species of Anomalobuthus known to occur south of the impressive Kopet Dagh mountain 2. The pioneering observations of Pavlovsky (1916b) range, the natural boundary between Turkmenistan and in Dzhulek (now Baigakum) remain the only data on this Iran (Fig. 141). species' biology and behavior. DISTRIBUTION (Fig. 141). Sands of the Dasht-e-Lut 3. The Baigakum specimens of this new species were Desert, in eastern Iran. included (under A. rickmersi) in a DNA phylo- geographic study by Graham et al. (2012), which ECOLOGY. See Teruel et al. (2014). demonstrated a derived position of this taxon within the genus. NOTES. See Teruel et al. (2014).

Anomalobuthus talebii Teruel, Kovařík, Anomalobuthus zarudnyi (Birula, 1911), comb. n. Navidpour et Fet, 2014 Figure 141 Figures 90–106, 141. Tables 3–5 Psammobuthus zarudnyi Birula, 1911a: 69–74; Birula, Anomalobuthus talebii Teruel, Kovařík, Navidpour & 1911b: 172; Birula, 1917: 131–132, 150, 166; Fet, Fet, 2014: 1–10, figs. 1–24, tabs. 1–2. 1989: 119; Fet & Lowe, 2000: 214 (includes full list of references before 1998); Capes & Fet, 2001: HOLOTYPE. ♀ (FKCP). Iran, South Khorasan Prov- 300–301; Fet et al., 2001: 184. ince, Hemmatabad (=Hemmatābād) Desert, 33°20'49. 45"N 60°25'56.86"E, close to the Afghanistan border, 17 LECTOTYPE ♂ (ZISP 615, herein designated). April 2013, leg. A. Talebi Gol. Uzbekistan, Andijon Province, Mingbulak, 40°51'56"N 71°39'55"E, 13 [26] August 1909, leg. N. Zarudny. DIAGNOSIS (updated, based on one adult female). Adult size 35 mm. Coloration yellow, with irregular PARALECTOTYPE. ♀ (ZISP 616). Tajikistan, Khu- blackish spots on carapace, tergites, pedipalps, legs, and jand Province, Kairakkum, 40°15'39"N 69°47'57"E, 28– metasoma; tergites with two faint submedian dark 31 May [10–13 June] 1908, leg. N. Zarudny.

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 29

Figures 90–91: Anomalobuthus talebii, adult female holotype: full-body views, dorsal (90) and ventral (91).

DIAGNOSIS (updated, based on one adult pair). Adult segment V composed of sharply lobate denticles, dorsal size standard for the genus (male 29 mm, female 30 lateral and lateral supramedian carinae of segments I–IV mm). Coloration yellow, with irregular blackish spots with terminal denticle enlarged; intercarinal areas mostly and reticulations on carapace, tergites, pedipalps, legs, smooth, with granulation sparse ventrally and laterally, and metasoma; tergites with two conspicuous submedian of segment V dense ventrally and sparse laterally. Tel- dark stripes; metasomal segment V and telson reddish. son vesicle elongate oval. Pedipalp fingers with 10–11 principal rows of denticles and 8 internal accessory denticles. Pectines with 24/26 DESCRIPTION (adult male lectotype, translated and teeth in male, 20/20 in female. Tibial spur present in legs updated from Birula, 1911a). Coloration base pale to III–IV (weaker in female leg III). Metasoma with most brownish yellow. Pedipalp femur with dorsal surface carinae moderately developed; ventral lateral carinae of infuscate; patella with internal surface infuscate. Cara- 30 Euscorpius — 2018, No. 270

Figures 92–100: Anomalobuthus talebii, adult female holotype: chelicerae, carapace and tergites I–III, dorsal (92), sternopectinal region and sternites III–V, ventral (93), distal part of right legs I–IV, internal (94–97), metasoma and telson, lateral (98), dorsal (99) and ventral (100). pace with an anterior V-shaped, broad blackish spot margin. Tergites each with two large dark spots, that from median ocular tubercle through frontal margin, form a pair of broad lateral bands. Legs essentially im- plus three large, transverse blackish spots: one behind maculate, only with femur of II–III infuscate. Metasoma median ocular tubercle and two laterally along posterior not conspicuously bicolor, but becoming progressively

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 31

Figures 101–106: Anomalobuthus talebii, adult female holotype: right pedipalp trochanter and femur, dorsal (101), right pedipalp patella, dorsal (102) and external (103), right pedipalp chela, dorsal (104) and external (105), right pedipalp movable finger, dorsal (106). Black dots depict trichobothria. darker distally, with a diffuse annulated appearance: ba- numbering eight on both fingers, movable finger with sal part of each segment infuscate in the shape of dark one accessory denticle basal to the terminal denticle. rings, which become larger and reticulate both dorsally Carapace. Anterior margin straight to shallowly and ventrally. convex. Carination essentially absent: the only carinae Pedipalps. Relatively short but very slender. Femur present are the superciliaries, which are strongly granu- with carinae granulose; intercarinal tegument smooth lose. Tegument very finely and densely granulose, with and glossy. Patella with carinae vestigial, smooth; inter- coarser granules scattered all over. carinal tegument smooth and glossy. Chela elongate and Sternum. Standard for the genus, relatively small slender; manus somewhat wider than patella (ratio 1.50), and wide. somewhat flattened (1.07 times longer than wide, 1.67 Genital operculum. Each half roundly subtri- times wider than deep), without carinae; intercarinal angular in shape. tegument smooth and glossy; fingers long (movable Pectines. Standard-sized for the genus: very long, finger 2.00 times longer than underhand), only subtly extending to posterior margin of sternite IV, sub- curved and with 10–11 principal rows of denticles (the rectangular. Tooth count 24/26. three basalmost rows are poorly defined), basal Legs. Very slender, with all carinae weak and sub- lobe/notch combination absent, external accessory den- granulose to subcostate; intercarinal tegument smooth ticles absent, internal accessory denticles large and and glossy. Tibial spurs well-developed on legs III–IV.

32 Euscorpius — 2018, No. 270

Figures 107–119: Anomalobuthus spp., from ZISP historical collection: adult male topotypes of A. pavlovskyi sp. n. (107– 112), full-body dorsal views (107, 110), labels (108, 111) and bottles (109, 112); adult male (113) and adult female (114) A. pavlovskyi sp. n. from Kunya-Urgench, Turkmenistan, full-body dorsal views (113, 114), labels (115) and bottle (116); adult male A. rickmersi from Tejen, Turkmenistan (117–119), full-body dorsal view (117), labels (118) and bottle (119). Photographs courtesy Alexander Koval.

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 33

Figures 120–125: Anomalobuthus krivochatskyi sp. n., adult male paratopotype: SEM views of chelicerae, dorsal and ventral (120), metasoma V and telson, lateral (121); anterior half of carapace, dorsal (122); close-up of trichobothria (123); left leg IV basitarsus and telotarsus, ventrointernal (124); left leg IV basitarsus and telotarsus, external (125). Images courtesy Jan Štundl.

34 Euscorpius — 2018, No. 270

A. lowei sp. n. A. krivochatskyi sp. n. A. pavlovskyi sp. n. A. rickmersi A. talebii Ratios ♀ HT ♂♂ HT, PT ♀♀ PT ♂ HT ♀♀ PT ♂ HT ♀♀ ♀ HT Pedipalp chela (L/W) 6.41 6.20–6.86 5.86–5.87 6.80 5.83–6.90 6.20 5.58–6.20 6.29 Pedipalp chela (L) / movable finger (L) 1.43 1.38–1.45 1.41–1.42 1.42 1.39–1.40 1.41 1.31–1.39 1.44 Pedipalp movable finger (L) / manus (L) 2.31 2.20–2.61 2.42–2.50 2.40 2.50–2.55 2.44 2.55–3.18 2.26 Metasoma + Telson (L) / Carapace (L) 5.40 6.26–6.37 6.12–6.13 7.14 5.68–5.77 7.67 6.05–6.38 6.34 Metasomal segment I (L/W) 1.02 1.26–1.35 1.09–1.23 1.31 1.22–1.31 1.74 1.43–1.44 1.30 Metasomal segment II (L/W) 1.35 1.76–1.77 1.50–1.62 1.75 1.74–1.83 2.13 1.84–1.96 1.82 Metasomal segment III (L/W) 1.39 1.83–1.86 1.64–1.71 1.77 1.85–1.86 2.20 2.04–2.08 1.88 Metasomal segment IV (L/W) 1.75 2.11–2.14 1.83–1.89 2.23 2.05–2.15 2.56 2.30–2.31 2.23 Metasomal segment V (L/W) 1.85 2.32–2.33 2.00–2.11 2.38 2.16–2.31 2.71 2.42–2.50 2.34 Telson vesicle (L/W) 1.47 1.92–2.15 1.75–1.80 1.89 1.66–1.89 2.50 1.89–2.00 2.18 Telson vesicle (L/D) 1.66 1.77–2.18 1.91–1.93 2.12 2.08–2.12 2.39 1.89–2.17 2.14

Table 4: Comparison between five species of Anomalobuthus based upon selected morphometric ratios. Abbreviations: length (L), width (W), depth (D), holotype (HT), paratypes (PT).

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 35

Figures 126–133: Anomalobuthus spp., SEM views of right pedipalp of adult male A. rickmersi from Repetek (126–127) and adult male paratopotype A. krivochatskyi sp. n. (128–133): chela (126, 128), close-up of manus (127, 129), three ventrointernal magnified views of mid part of both fingers in consecutive figures (130–132, movable finger on top), ventrointernal close-up of fingertips (133, movable finger on top). See trichobothria highlighted in red.

Mesosoma. Tergites very finely and densely gran- carina is weak and the submedian carinae are poorly ulose, with subtle vestiges or coarser granules scattered; defined by 2–3 coarse granules only. Sternites III–VI I–VI irregularly tricarinate: the median longitudinal smooth and glossy; VII with two pairs of carinae: the 36 Euscorpius — 2018, No. 270

Figures 134–140: Anomalobuthus spp., SEM views showing size variation of leg IV tibial spur (highlighted in red): large in paratopotype female A. krivochatskyi sp. n. (134), moderately large in paratype female A. krivochatskyi sp. n. from Bukhara (135), small in paratopotype female A. krivochatskyi sp. n. (136), absent in male A. rickmersi from Repetek (137), close-up of a medium-sized leg IV tibial spur of a paratopotype female A. krivochatskyi sp. n. from Zarafshan (138), see spinulate and striate sculpture. Anomalobuthus krivochatskyi sp. n., adult female paratopotype: SEM views of right pecten and adjacent parts of leg coxae and sternite III (139), peg sensilla of tooth from same pecten (140).

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 37

Species Sex N Pectinal Teeth Ave. SD 19 20 21 22 23 24 25 26 27 ♂♂ 8 3 1 3 1 23.25 ± 1.09 A. krivochatskyi sp. n. ♀♀ 16 2 7 4 3 20.50 ± 0.94 A. lowei sp. n. ♀♀ 2 1 1 20.50 ± 0.50 ♂♂ 15 2 6 5 2 25.47 ± 0.88 A. pavlovskyi sp. n. ♀♀ 12 8 4 22.33 ± 0.47 ♂♂ 22 4 4 7 6 1 24.82 ± 1.15 A. rickmersi sp. n. ♀♀ 9 2 1 3 3 21.78 ± 1.13 A. talebii ♀♀ 2 2 26.00 ± 0.00 ♂♂ 2 1 1 25.00 ± 1.00 A. zarudnyi comb. n. ♀♀ 2 2 20.00 ± 0.00

Table 5: Pectinal tooth count variation in all known species of Anomalobuthus. Abbreviations: number of pectines (N), average (Ave.), standard deviation (SD). This includes counts taken from sources with precise and reliable geographic data (Birula, 1905, 1910).

submedians are smooth and the laterals are granulose, VARIATION. Unknown, only one adult of each sex intercarinal tegument very finely and densely granulose have been collected. except smooth medially. Metasoma. Moderately elongated and slightly nar- COMPARISON. According to Birula (1911a), the ex- rower distally; with 10/8/8/8/5 complete to almost com- tensively dark-patterned coloration and the strong plete, serrate carinae: dorsal laterals very strong on I, development of ventral lateral and ventral submedian strong on II–III with terminal denticles enlarged on the carinae on metasomal segments II–III, both seem to latter, moderate on IV, absent on V; lateral supra- distinguish this species clearly from all its other medians very strong on I, irregular on II–III with congeners. Moreover, it is the only species of terminal denticles enlarged on the latter, weak on IV, Anomalobuthus known to occur in the sands of the smooth on V; lateral inframedians very strong on I, Ferghana Valley. undefined to absent on II–V; ventral laterals strong on I, strong on II–III, obsolete on IV, strong on V, where DISTRIBUTION (Fig. 141). Isolate sands of the Ferg- become progressively stronger and somewhat flared hana Valley in the border region between Uzbekistan distally, formed by sharp, subequal denticles; ventral and Tajikistan. submedians very strong on I, strong on II–III, obsolete on IV, absent on V; ventral median absent on I–IV, NOTES. finely denticulate and distally bifurcate on V. Inter- 1. The only known two specimens of this species were carinal tegument smooth and glossy, with many granules collected in 1908 and 1909 by the famous Russian of different sizes scattered all over lateral and ventral ornithologist and traveler Nikolay Alekseevich Zarudny surfaces, which become progressively denser and coar- (1859-1919) who also brought numerous scorpions for ser towards segment V. Dorsal furrow moderately deep A. Birula's ZISP collection, including many new species on all segments. Setation sparse over carinae. from the previously unexplored areas of Iran. Telson. Vesicle elongate oval (2.00 times longer 2. The unavailability of the types and lack of than wide, 0.77 times wider than deep), tegument additional specimens of A. zarudnyi comb. n. prevents smooth and glossy, with coarse granules arranged into us from doing a deeper analysis of its true taxonomic longitudinal carinae. Subaculear tubercle absent. Acu- status within the genus. Until any counterevidence leus shorter than vesicle. appears, it seems more appropriate to retain it as a valid species. Moreover, two points seem to support the pre- FEMALE (paralectotype, data extracted from Birula, sent assumption: the strikingly contrasting coloration 1911a). Very similar to male, sexual dimorphism evident (Birula, 1911a) that does not match any of the pop- by: 1) size slightly larger; 2) metasoma somewhat less ulations of Anomalobuthus studied during this revision, slender; 3) tergites I–VI with median carina more and its isolated distribution inside the Ferghana Valley strongly and regularly developed; 4) pectines with con- (Fig. 137). sistently lower tooth count (20/20); 5) tibial spur highly 3. In the last 100 years, the Ferghana Valley sands reduced on leg III. have all but disappeared due to irrigation. On 18-20 May 38 Euscorpius — 2018, No. 270

2002, our field expedition (VF and A.V. Gromov) Discussion visited the remaining sand massifs of the modern Ferghana Province, Uzbekistan (Kairakkum Sands in Morphology Besharyk District, 40.4735°N 70.4503°E, and “Kara- kalpak Steppe” in Yazyavan District, 40.6580°N Trichobothria. The trichobothria of the genus 71.5072°E). We did not find A. zarudnyi, while psamm- Anomalobuthus present a very peculiar problem. Even ophilic Mesobuthus gorelovi Fet et al., 2018 was abun- though the trichobothrial pattern is clearly A-β, some dant and active. trichobothria are either absent or very reduced and

Key to the species of Anomalobuthus difficult to distinguish from the surrounding setae under ordinary white/UV light microscopy, e.g., femoral d2 Kraepelin, 1900 and chelal Et, V , and esb. This is because most ordinary 1 1. Coloration with the dark pattern very dense all over, pedipalpal setae are hypertrophied, having a very large tergites with two conspicuous submedian stripes …..… and heavily socketed base and a thick shaft, too closely …………………………………………..…. A. zarudnyi resembling true trichobothria which also have shaft - Coloration with the dark pattern inconspicuous to es- thicker than usual (Figs. 72–74, 122–129). Despite this, sentially absent on carapace, legs and metasoma (except its current placement in the “Buthus group” by Fet et al for segment V blackish to reddish), tergites immaculate (2005) stands correct, based on two important diagnostic to at most with two faint submedian stripes ………….. 2 characters: trichobothrial pattern beta, with patellar d3 trichobothrium located internal to dorsal median carina. Tibial spurs. Fet et al. (2005) reviewed the impor- 2. Tergites with two faint submedian dark stripes. Fe- tance of tibial spur on legs III–IV as a character in male (single sex known) with 26 pectinal teeth ………. scorpion systematics, in particular in buthoid scorpions. ………………………………………..………. A. talebii In parvorder Buthida (sensu Soleglad & Fet, 2003b), - Tergites without submedian dark stripes. Female with tibial spurs are completely absent (lost) in New World 19–23 pectinal teeth ………………………………….. 3 genera (a major synapomorphy) but their presence is variable within the Old World members (Fet et al., 2005, 3. Habitus very robust in female (single sex known), see Table 1). the following ratios: metasoma + telson length/carapace In his original diagnosis of the male holotype of A. length = 5.40, metasomal segment I length/width = 1.02, rickmersi, Kraepelin (1899) reported that the tibial spur metasomal segment V length/width = 1.85, telson ves- was absent on legs III, and present but reduced (to a very icle length/depth = 1.66 ………..………. A. lowei sp. n. small size, represented by a sinuous remnant) on legs IV. - Habitus remarkably slenderer in female, see the fol- His important text is translated below: lowing ratios: metasoma + telson length/carapace length = 5.68–6.38, metasomal segment I length/width = 1.09– “Recently I got from Professor O. Schneider a small 1.44, metasomal segment V length/width = 2.00–2.50, scorpion collected in the Buchara by Mr. W. telson vesicle length/depth = 1.89–2.17 …………..….. 4 Rickmer Rickmers, which unfortunately contradicts with classification of Buthidae as established by 4. Adult metasoma with intercarinal spaces much more myself, into the two subfamilies of Buthinae and densely granulose in female than in male; basal third of Centrurinae because, although an undoubtedly gen- uine Buthine, a character feature of this group, a segment V yellowish red ………….. A. pavlovskyi sp. n. tibial spur of the third and fourth pairs of legs, in this - Adult metasoma with intercarinal spaces either smooth scorpion is developed only at the fourth leg pair as a and glossy in female, or more densely granulose in male; very tiny rudiment (Fig. 4), being completely absent segment V entirely reddish to blackish, at most only on the third pair of legs. Since the structure of the subtly paler at its extreme base ……………………….. 5 rows on the cutting edge of pedipalp finger also does not match with any of the existing genera, I decided 5. Adult male habitus very slender: metasoma + telson to establish a new genus Anomalobuthus, which I length/carapace length ratio = 7.67. Adult metasoma must attribute, despite its almost completely dis- appearing tibial spur, to the subfamily of Buthinae, with ventral and lateral intercarinal spaces sparsely as discussed below in more detail.” granulose on segment IV and very densely granulose on V, especially in male; segment V reddish in adult male Birula (1905) following Kraepelin made a connec- ……………...…………………………….. A. rickmersi tion between this reduction and the absence of tibial - Adult male habitus less slender: metasoma + telson spurs in the New World buthids, thus questioning the length/carapace length ratio = 6.26–6.37. Adult meta- relationship of Anomalobuthus with other Old World soma with intercarinal spaces almost smooth in both genera. Birula (1905: 449–450) who was the next to sexes; segment V blackish in adult male ………….. A. study Anomalobuthus, had only two specimens collected krivochatskyi sp. n. in the modern Turkmenistan. He observed that a male

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 39

Figure 141: Distribution of the genus Anomalobuthus: A. krivochatskyi sp. n. (1), A. lowei sp. n. (2), A. pavlovskyi sp. n. (3), A. rickmersi (4), A. talebii (5), A. zarudnyi comb. n. (6), and literature records whose identification is pending (black squares). Data compiled from specimens studied herein and the following supplementary sources: Rickmer Rickmers (1913), Fet (1989, 1994), Gromov & Kopdykbaev (1994), Gromov (2006), Mityaev et al. (2008), and Graham et al. (2012). from Tejen (ZISP 809, clearly diagnosable as a male by mentioned tibial spur reduction in Anomalobuthus. having 25 pectinal teeth, see our Figs. 113–115) had Sissom (1990: 96) stated that in this genus tibial spurs developed tibial spurs only on legs IV, while a female on legs III are absent, and on legs IV are “sometimes from Repetek (ZISP 808, 22 pectinal teeth) completely vestigial”. Note that Sissom (1990) only studied Turk- lacked tibial spurs. Later, Birula (1911a: 73) in des- menistan specimens from Repetek, supplied by V. Fet. cribing the genus Psammobuthus Birula, 1911, from Later, Capes & Fet (2001) and Fet et al. (2001) stated Ferghana (Uzbekistan) (which we here synonymize with that females of Anomalobuthus lack tibial spurs while Anomalobuthus) noted that it had well-developed tibial males have them “occasionally present, but small, on leg spurs on legs III and IV in males but rudimentary ones IV.” on legs III in females; and stated that “females in Our current study revealed a complex picture of Anomalobuthus have no spurs on legs III and IV but interspecific, intraspecific, and intrapopulational vari- males only have rudimentary tibial spurs on legs IV.” ation in loss or reduction of tibial spurs across all six This was interpreted as a sexual dimorphism. Birula species of the genus Anomalobuthus (Figs. 130–134). (1911b: 171) reported two more males from Baigakum This variation includes several stages of reduction, (Kazakhstan), collected by D. Glazunov, but did not marked fluctuating asymmetry (presence on either left or comment on their tibial spurs. Males in Baigakum (our right side), and possible sexual dimorphism. The tibial A. pavlovskyi sp. n.), as we now know, have spurs in spur was usually absent in A. rickmersi from Turk- different degrees of development. menistan (except the male from Tejen has it on both legs All subsequent genus-level keys of Buthidae IV), and is absent in the only known female of A. talebii (Werner, 1934; Stahnke, 1972, fig. 3; Sissom, 1990) from Iran. In A. krivochatskyi sp. n., the tibial spur was 40 Euscorpius — 2018, No. 270 always absent in males, but in females it was present and we suggest that it represents a derived character always on both legs IV, and sometimes also in leg III but state” (Navidpour & Lowe, 2009). only on one side. In A. pavlovskyi sp. n., the tibial spur For another psammophile genus from the Arabian development was highly variable among Baigakum Peninsula, Vachoniolus, Lowe (2010) noted that, in V. specimens: it was either entirely absent from all legs gallagheri, “tibial spurs were either present or absent, (one male), or present asymmetrically on a single leg IV and were absent more frequently on leg III than leg IV; (three males including holotype, two females), or present of 30 leg III tibiae, spurs were present in 19 (63.3 %), asymmetrically on each leg III–IV of the same side (left, and absent in 11 (36.7 %); of 29 leg IV tibiae, spurs one female), or, finally, present on both legs IV (two were present in 25 (86.2 %), and absent in 4 (13.8 %).” males, one female); but in all cases where it was present, The same trend was observed by Lowe (2010) in the it was small to vestigial. In the only known female of A. congeneric V. globimanus: “Tibial spurs were either pre- lowei sp. n., tibial spur is absent from both legs III and sent or absent, or vestigial (< 20% of the length of fully right leg IV, but a moderate-sized spur is found on left formed spur), and were lost with similarly high leg IV. Finally, in A. zarudnyi sp. n., the only known frequencies on both legs III and IV: of 78 intact leg III male has tibial spurs well-developed on legs III–IV but tibiae, spurs were present in 39 (50.0 %), and absent or in the only known female, they are highly reduced on vestigial in 39 (50.0 %); of 77 intact leg IV tibiae, spurs legs III. These patterns suggest a clear trend within were present in 43 (55.8 %), and absent or vestigial in 34 Anomalobuthus for developmental instability in exhib- (44.2 %).” iting variable reduction or loss of an important morpho- Importantly, already the pilot DNA-based phylo- logical character. geny of Fet et al. (2003) for 17 genera of Buthidae Quite possibly, this trend is related to psammophilic suggested the polyphyletic origin of psammophily adaptation of Anomalobuthus since it is also observed in among these genera; full or partial tibial spur loss is other Old World sand desert scorpions. Various observed independently in at least three independent modifications of leg structures in psammophile buthids lineages which include (a) Anomalobuthus, (b) Lio- have been amply documented (Fet et al., 1998). Mono- buthus, and (c) Vachoniolus + Apistobuthus. We suspect typic genera Liobuthus Birula, 1898 and Pectinibuthus therefore that this trend is an example of parallelism Fet, 1984 (Central Asia) and Plesiobuthus Pocock, 1900 (convergent evolution) under selection toward psammo- (Pakistan) completely lack tibial spurs (Capes & Fet, phily. 2001; Fet et al., 2001, 2005). Tibial spurs are often reduced or missing, with asymmetry exhibited, in two Biogeography Middle Eastern psammophile buthid genera, Apisto- buthus Finnegan, 1932 and Vachoniolus Levy et al., The dynamic biogeographic history of Central 1973, which have been recently studied in detail. Asian sand deserts is well-known, and likely had a pro- Graeme Lowe (pers. comm.) kindly summarized for us found impact on the diversification of psammophilic the available data. Already in her original description of scorpions, including remarkable endemic genera Ano- Apistobuthus pterygocercus, Finnegan (1932) stated: malobuthus and Liobuthus. The fauna, flora and geology “Tibial spurs absent on third pair of legs and present as a of Kazakhstan, Uzbekistan, and Turkmenistan have been small weak spine on only one of the fourth pair of legs thoroughly studied but most of the literature was in two of the three specimens collected (text-fig. 8), published in Russian, thus remaining largely inaccess- absent from all the legs of the third specimen.” Such ible to the global research community. The most com- asymmetric reduction or loss in this species was also prehensive biogeographic review was generated for reported by Vachon (1980). Navidpour & Lowe (2009) Coleoptera (Kryzhanovsky, 1965), while the remaining found that tibial spur loss or degeneration (to a very data on the fauna and biogeography of Central Asia are small vestigial spur ca 20% the length of a fully scattered in Russian journals as dozens of papers cover- developed spur) was much more common in A. ing specific taxa and regions. After the demise of the pterygocercus than in the congeneric A. susanae. “In a USSR in 1991, the former Soviet Central Asia was sample of n = 20 adults of A. susanae: 39 intact leg III fragmented politically into five independent countries tibia included 30/39 spurs present, 9/39 (23%) spurs lost; (Kazakhstan, Kyrgyzstan, Uzbekistan, Tajikistan, and 38 intact leg IV tibiae included 34/38 spurs present, 4/39 Turkmenistan). Since then, the rate and quality of (10%) spurs lost. In contrast, in a sample of n = 31 zoological research from the region has decreased adults of A. pterygocercus: 60 intact leg III tibiae markedly. Of these five countries, only a comprehensive included 15/60 spurs present, 45/60 (75%) spurs lost; 60 review of the ecology and biogeography of Turk- intact leg IV tibiae included 36/60 spurs present, 24/60 menistan has been published in English (Fet & Ata- (40%) spurs lost. The much higher frequency of muradov, 1994). Our research group and its collab- degeneration or loss of tibial spurs in A. pterygocercus orators were the first to use molecular phylogenetics correlates to the ultrapsammophile habit of this species, (based on mitochondrial DNA sequence data) to study

Teruel, Kovařík & Fet: Revision of Genus Anomalobuthus 41 the biogeography of scorpions from the Central Asian Graham et al. (2012) confirms that both great waterways deserts (Gantenbein et al., 2003; Graham et al., 2012; of Central Asia, especially Amudarya and (in part) Fet et al., 2018). Syrdarya, acted as biogeographic boundaries in vicariant As with many animal and plant groups of Central speciation of Anomalobuthus. They indicated also that Asia, speciation is clearly connected to the region's “the levels of divergence within Anomalobuthus, Lio- geomorphology; such as landscape fragmentation driven buthus, and Mesobuthus are somewhat deep for by mountain uplift and substrate modifications during intraspecific studies.” Indeed, the most recent revision of the formation of sand and clay deserts. Furthermore, the “Mesobuthus caucasicus complex” (Fet et al., 2018) enclaves of sand desert within mountain valleys (such as revealed existence of diverse species across Central the type locality of A. rickmersi in Tajikistan) con- Asia. tributed to local evolution of psammophilic fauna, with The southern and southeastern Turkmenistan narrow endemics found at considerable altitude. In Cen- populations (Karakum Desert), which morphologically tral Asia, geological data emphasize two major types of all belong to A. rickmersi, are clearly separated by the late Cenozoic paleogeographical changes, mountain Amudarya River from the Uzbekistan/southern Kaz- uplift and eustatic changes of the ancient Caspian Sea (a akhstan populations (Kizylkum Desert), which are now remnant of the Tethys Sea). The great alluvial deserts, demonstrated to belong to a morphologically distinct A. primarily the Karakum and the Kizylkum, have been krivochatskyi sp. n. A more complex picture emerges formed as a result of deposits by the Amudarya and regarding the vicariant distribution of A. krivochatskyi Syrdarya Rivers (reviewed in Graham et al., 2012). sp. n., limited to the southern Kizylkum (40-41°N) and The largely psammophilic Mesobuthus gorelovi Fet A. pavlovskyi sp. n., found in the sands of the Aral Basin et al., 2018., which spans most of the deserts of Central (42-44°N), from extreme northern Turkmenistan (Kunya Asia, shows significant phylogeographic structure, with Urgench) to south-central Kazakhstan (Baigakum, or specimens from Turkmenistan forming a clade distinct Dzhulek on Syrdarya River); in addition, we describe the from populations in Uzbekistan and Kazakhstan. M. northernmost Kazakhstan species A. lowei sp. n. from gorelovi is one of the most common scorpion species in the Ili River valley (Kapchagay). sp. n.. The DNA data Central Asia, found widespread in the lowland deserts of Graham et al. (2012) indicate that A. krivochatskyi sp. and exhibiting psammophilic adaptations such as sand n. forms a sister clade to A. pavlovskyi sp.n, as compared combs. The distribution of M. gorelovi was likely af- to an outgroup A. rickmersi. An approximate divergence fected by transgressions of the Caspian Sea, especially in time between two new species is estimated from 16S Pliocene and further in Pleistocene, when it was divided mtDNA data (M. R. Graham, pers. comm.) between 5.34 by the Amudarya River, as found in other co-distributed and 1.62 Ma, with a mean estimate of 3.41 Ma (Plio- buthids (Graham et al., 2012). Divergence within M. cene), which corresponds well to the above listed gorelovi was estimated to have occurred in this Mesobuthus divergence estimates. Early to Middle timeframe (5.2 – 1.4 Ma; mean = 3.1 Ma). The main Pliocene has been the time of diversification in many clades within M. gorelovi occur on either side the taxa in the deserts and mountains of Central Asia, Amudarya River, consistent with the hypothesis of followed by the Caspian Sea transgressions and cooling Pliocene vicariance. Another psammophilic species, of the climate in the late Pliocene (Atamuradov, 1994; Mesobuthus elenae Fet et al. 2018, was found isolated in Kazenas & Baishashov, 1999). the Amudarya Valley of southwestern Tajikistan and Allopatric distribution of species-level, uniquely Uzbekistan, broadly overlapping with the only known adapted psammophile endemics in the alluvial sands of Tajikistan population of A. rickmersi (Baljuvon). The the Karakum, Kizylkum, and the Aral Basin has been divergence of M. elenae from its sister species M. par- studied in many examples of invertebrate and vertebrate thorum is also dated to the Pliocene (4.9 – 1.6 Ma; mean fauna (Kryzhanovsky, 1965). As one of the most recent = 3.1 Ma). studies, we can quote a comprehensive molecular and Graham et al. (2012) have analyzed mtDNA mar- morphological work on the psammophilic toad agama kers in several specimens of Anomalobuthus, then Phrynocephalus mystaceus (Agamidae) (Solovyova et placed under “A. rickmersi”. As we demonstrate above al., 2018), which sampled many of the same desert on the basis of morphology, the material used by localities as our work on scorpions. We are glad to Graham et al. (2012) should be classified as three report that Anomalobuthus follows the classical pattern different species, namely A. rickmersi (Turkmenistan), of vicariant speciation in these great sand deserts of A. krivochatskyi sp. n. (Uzbekistan) and A. pavlovskyi Central Asia. sp. n. (Kazakhstan). Although this molecular study did not include A. lowei sp. n., A. talebii Teruel et al., 2014, Acknowledgments and A. zarudnyi (Birula, 1911), comb. n., its conclusions are very important in addressing the endemic speciation The 2002 field mission to Central Asia (Kazakhstan, of Anomalobuthus. This first DNA-based analysis of Turkmenistan, and Uzbekistan) that provided many of 42 Euscorpius — 2018, No. 270 the studied specimens was supported by the National gaucho (Scorpiones, Buthidae), a new species from Geographic Society (USA) Research and Exploration southern Brazil, with an update about the generic Fund (grant 7001–0001 to VF). Alexander Gromov pro- diagnosis. The Journal of Arachnology, 36: 491– vided a great help during the field work across Central 501. Asia in March–May 2002. Collection permits were granted by the ministries of natural resources of ARMAS, L. F. DE, R. TERUEL & F. KOVAŘÍK. 2011. Kazakhstan, Turkmenistan, and Uzbekistan. We are Redescription of granosus (Thorell, grateful to Dzhamshid Dzhuraev, Aliya Gromova, 1876) and identity of Centrurus granosus simplex Gochmyrat Gutlyev, Alexander and Elena Kreuzberg, Thorell, 1876 (Scorpiones: Buthidae). Euscorpius, Viktor Lukarevsky, Sergei Morozov, Shukhrat Shak- 127: 1–11. hnazarov, and Alexander Tarabrin for their hospitality and help in field logistics. ATAMURADOV, KH. I. 1994. Paleogeography of We also thank everyone who helped to obtain and Turkmenistan. Pp. 49–64 in: Fet, V. & Kh. I. describe rare specimens in field and from the museum Atamuradov (eds.). Biogeography and Ecology of collections over many years, including but not limited Turkmenistan. Kluwer Academic Publishers, Dord- to: Yuri Balashov, Matt Braunwalder, Michelle Capes, recht. Hieronymus Dastych, Andrei Feodorov, Yuri Gala- ktionov, Yuri Gorelov, Alexander Gromov, Jürgen Gru- ber, Danilo Harms, Christoph Hörweg, Ragnar Kinzel- BIRULA, A. A. 1904. Miscellanea Scorpiologica. VII. bach, Viktor Krivochatsky, Pavel Kučera, Gennady Synopsis der russischen Skorpione. Annuaire du Kuznetsov, Yuri Marusik, Gleb Medvedev, Kirill Musée Zoologique de l'Académie Impériale des Mikhailov, Vladimir Ovtsharenko, W. David Sissom, Sciences de St. Pétersbourg, 9: 28–38. Oraz Soyunov, Verena Stagl, and Andreas Wessel, and Sergei Zonstein. BIRULA, A. A. 1905. 4. Skorpiologische Beiträge, 1.-3. We are grateful to Graeme Lowe for his great help Microbuthus littoralis (Pavesi), Anomalobuthus in taking exquisite images of the rare A. lowei sp. n. We rickmersi Kraepelin und Buthus zarudnianus n. especially thank Alexander Koval, Andrei Ovchinnikov, nom. Zoologischer Anzeiger, 29(1-4): 445–450. and Anastasia Ovchinnikova (St. Petersburg, Russia) for taking the images of the ZISP specimens, and Kirill BIRULA, A.A. 1911a. Skorpiologische Beiträge. 7.-8. Mikhailov, Elena Temereva, and Roman Nazarov Psammobuthus g. n. Zoologischer Anzeiger, 37(3- (Moscow, Russia) for the images of the ZMMSU 4): 69–74. specimens; and Jan Štundl for several scanning electron microscope (SEM) images (Figs. 116–121). We thank BIRULA, A. A. 1911b. Miscellanea Scorpiologica. IX. Matthew R. Graham, Viktoria Oláh-Hemmings, and Ein Beitrag zur Kenntnisse der Skorpionenfauna des Joshua Greenwood for the DNA analysis that allowed Russischen Reiches und der angrenzenden Länder. the first insight into Anomalobuthus phylogeny and Annuaire du Musée Zoologique de l'Académie biogeography (Graham et al., 2012). David Neff, Impériale des Sciences de St. Petersburg, 16: 161– Michael Brewer, Matthew R. Graham, Jacqueline Ais- 179. ling (Webb), Kelly Anne Daniel, and Kelsey Longe greatly helped VF with SEM imaging (Figs. 122–136). Shakhrokh Navidpour brought to our attention (Teruel et [BIRULA, A. A.] BYALYNITSKII-BIRULYA, A. A. al., 2014) the only known Iranian species, which was 1917. 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